crecimiento, rendimiento y contenido de azúcares a diferente edad fisiológica del tubérculo de...

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Growth, yield and sugar content of potato tubers at different physiological ages Crecimiento, rendimiento y contenido d e azúcares a di ferente edad fisiológica del tubérculo de papa Sigfrido David Morales-Fernández *; Rafael Mora-Aguilar ; Yolanda Salinas-Moreno ;  Juan Enrique Rodríguez-Pérez ; María Teresa Colinas-León ; Héctor Lozoya-Saldaña  Universidad Tecnológica de la Selva. Departamento de In vestigación y Desarrollo. Carretera Ocosingo - Altamirano km ., Ocosingo, Chiapas, C. P. , MÉXICO. Correo-e: moralesuno@hotmail.com, teléfono: (*Autor para correspondencia). Instituto de Horticultura. Departamento de Fitotecnia. Universidad Autónoma Chapingo. Carretera México - Texcoco km ., Chapingo, Estado de México. C. P. , MÉXICO. Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias. Centro - Altos de Jalisco. Carretera Tepatitlán - Lagos de Moreno km , colonia Rancho las Cruces, Tepatitlán de Morelos, Jalisco, C.P. , MÉXICO.  Abstrac t P otato tuber sugar content is an indicator of management conditions during crop growth, since it affects t he degree of maturity, quality and sprouting. In th is study, potato growth and yield under field conditions were assessed.  Soluble sugar content in tubers at different physiological ages was determined, as was the effect these sugars have on seed-tuber sprouting during storage. To this end, four tuber samplings were carried out in four potato cultivars: one during tuber dormancy and three at different physiological ages. Determinations were made using high-performance liquid chromatography (HPLC). The variety Alpha had the longest biol ogical cycle among all genotypes, and its y ield was similar to that of the varieties Gigant and  Vivald i ( g) , which had yields % greater th an that of the var iety Atla ntic. Potato tubers w ith the physiological age of apical dominance ( days after harvest, DAH), multiple sprouting ( DAH) and senescence ( DAH) had the highest sucrose, glucose and fructose contents (. to . mg·g - ), and the variety Vivaldi had the highest amount of sucrose and reducing and total sugars. The length and dry weight of the potato sprout showed a high positive and significant association with the content of reducing and total sugars. Resumen E l contenido de azúcares en el tubérculo de papa permite conocer las condiciones de manejo durante el crecimiento del cultivo; el grado de madurez, la calidad y su efecto en la brotación. En la presente investigación se evaluó el crecimiento y rendimiento de papa en condiciones de campo. Se determinó el contenido de azúca res solubles de tubérculos en diferentes edades fisiológicas, así como el efecto que tienen en la brotación de semilla-tubérculo durante su almacenamiento. Para ello, en cuatro cultivares de papa se realizaron cuatro muestreos de tubérculo: uno durante el reposo y tres en diferentes edades fisiológicas. Las determinaciones se realizaron mediante cromatografía líquida de alta resolución (HPLC). La variedad Alpha tuvo el ciclo biológico más largo entre todos los genotipos, y su rendimiento fue similar al de Gigant y  Vivaldi ( g) quienes resultaron % mayor que Atlantic. En los tubércu los de papa, con edad fisiológica de dominancia apical ( días después de la cosecha, DDC), brotación múltiple ( DDC) y senectud ( DDC), fue donde se presentaron los mayores contenidos de sacarosa, glucosa  y fructosa (. a . mg·g - ), y en los que Vivaldi registró la mayor cantidad de sacarosa, azúcares reductores y totales. La longitud y peso seco del brote de papa presentaron alta asociación positiva  y signif icativa con el contenido de azúcares reductores y totales. Received: June 27, 2014 / Accepted: July 1, 2015. Palabras clave: Solanum tuberosum L., fenología, peso de tubérculos, azúcares solubles, brotación. Keywords: Solanum tuberosum L., phenology, tuber weight, soluble sugars, sprouting. Please cite this article as follows (APA ):  Morales-Fernández, S. D., Mora-Aguilar, R., Salinas-Moreno, Y., Rodríguez- Pérez, J. E., Colinas-León, M. T., & Lozoya-Saldaña,  M. T. (). Growth, yield and sugar content of potato tubers at different physiological ages.  Revista Chapingo Serie Horticultura , (), . doi: ./r.rchsh.. . Scientific article www.chapingo.mx/revistas/horticultura

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7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 118

Growth yield and sugar content of potato tubersat different physiological agesCrecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del

tubeacuterculo de papa

Sigfrido David Morales-Fernaacutendez983089 Rafael Mora-Aguilar983090 Yolanda Salinas-Moreno983091 Juan Enrique Rodriacuteguez-Peacuterez983090 Mariacutea Teresa Colinas-Leoacuten983090 Heacutector Lozoya-Saldantildea983090

983089 Universidad Tecnoloacutegica de la Selva Departamento de Investigacioacuten y Desarrollo

Carretera Ocosingo - Altamirano km 983088983093 Ocosingo Chiapas C P 983090983097983097983093983088 MEacuteXICO

Correo-e moralesuno983089hotmailcom teleacutefono 983097983089983097983089983089983094983091983095983097983097 (Autor para correspondencia)983090Instituto de Horticultura Departamento de Fitotecnia Universidad Autoacutenoma Chapingo Carretera

Meacutexico - Texcoco km 983091983096983093 Chapingo Estado de Meacutexico C P 983093983094983090983091983088 MEacuteXICO983091Instituto Nacional de Investigaciones Forestales Agriacutecolas y Pecuarias Centro - Altos de Jalisco

Carretera Tepatitlaacuten - Lagos de Moreno km 983096 colonia Rancho las Cruces Tepatitlaacuten de Morelos Jalisco

CP 983092983095983094983088983088 MEacuteXICO

Abstract

Potato tuber sugar content is an indicator of management conditions during crop growth

since it affects the degree of maturity quality and sprouting In this study potato growth

and yield under field conditions were assessed Soluble sugar content in tubers at different

physiological ages was determined as was the effect these sugars have on seed-tuber sprouting

during storage To this end four tuber samplings were carried out in four potato cultivars one

during tuber dormancy and three at different physiological ages Determinations were made

using high-performance liquid chromatography (HPLC) The variety Alpha had the longest

biological cycle among all genotypes and its yield was similar to that of the varieties Gigant and

Vivaldi (983094983093983097 g) which had yields 983093983096 greater than that of the var iety Atlantic Potato tubers with

the physiological age of apical dominance (983089983088983097 days after harvest DAH) multiple sprouting (983089983093983091

DAH) and senescence (983090983091983095 DAH) had the highest sucrose glucose and fructose contents (983095983089983088 to

983095983096983097 mgmiddotg-983089) and the variety Vivaldi had the highest amount of sucrose and reducing and total

sugars The length and dry weight of the potato sprout showed a high positive and significant

association with the content of reducing and total sugars

Resumen

El contenido de azuacutecares en el tubeacuterculo de papa permite conocer las condiciones de

manejo durante el crecimiento del cultivo el grado de madurez la calidad y su efecto en la

brotacioacuten En la presente investigacioacuten se evaluoacute el crecimiento y rendimiento de papa en

condiciones de campo Se determinoacute el contenido de azuacutecares solubles de tubeacuterculos en diferentes

edades fisioloacutegicas asiacute como el efecto que tienen en la brotacioacuten de semilla-tubeacuterculo durante

su almacenamiento Para ello en cuatro cultivares de papa se realizaron cuatro muestreos de

tubeacuterculo uno durante el reposo y tres en diferentes edades fisioloacutegicas Las determinaciones se

realizaron mediante cromatografiacutea liacutequida de alta resolucioacuten (HPLC) La variedad Alpha tuvo elciclo bioloacutegico maacutes largo entre todos los genotipos y su rendimiento fue similar al de Gigant y

Vivaldi (983094983093983097 g) quienes resultaron 983093983096 mayor que Atlantic En los tubeacuterculos de papa con edad

fisioloacutegica de dominancia apical (983089983088983097 diacuteas despueacutes de la cosecha DDC) brotacioacuten muacuteltiple (983089983093983091

DDC) y senectud (983090983091983095 DDC) fue donde se presentaron los mayores contenidos de sacarosa glucosa

y fructosa (983095983089983088 a 983095983096983097 mgmiddotg-983089) y en los que Vivaldi registroacute la mayor cantidad de sacarosa azuacutecares

reductores y totales La longitud y peso seco del brote de papa presentaron alta asociacioacuten positiva

y significativa con el contenido de azuacutecares reductores y totales

Received June 27 2014 Accepted July 1 2015

Palabras clave Solanum

tuberosum L fenologiacutea pesode tubeacuterculos azuacutecares

solubles brotacioacuten

Keywords Solanum

tuberosum L phenologytuber weight soluble

sugars sprouting

Please cite this article as follows (APA 983094) Morales-Fernaacutendez S D Mora-Aguilar R Salinas-Moreno Y Rodriacuteguez-Peacuterez J E Colinas-Leoacuten M T amp Lozoya-Saldantildea M T (983090983088983089983093) Growth yield and sugar content of potato tubersat different physiological ages Revista Chapingo Serie Horticultura 983090983089(983090) 983089983090983097ndash983089983092983094 doi 983089983088983093983089983093983092rrchsh983090983088983089983092983088983094983088983091983089

Scientific article

wwwchapingomxrevistashorticultura

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 218

983089983091983088 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Introduction

The potato (Solanum tuberosum L) is the fourth mostimportant crop in the world (Sonnewald amp Sonnewald983090983088983089983092) Tuber formation depends on among other factorsthe availability of assimilates and the ability of tubersto accumulate them At the start of crop developmentphotoassimilates are mainly directed towards the

development of leaves stems and roots However thisbehavior changes with tuber initiation and growthbecause they increase the demand for assimilates (Wolf983089983097983097983091) and consequently the carbohydrates producedin the leaves throughout the crop cycle are translocatedto tuber structures

During tuber initiation starch content is low but thatof sugars is high since the rate of transport of sugarsproduced in the leaves exceeds the rate of conversion tostarch As a result when the tuber reaches physiologicalmaturity the maximum dry matter accumulation andminimum amount of sugars are obtained The latter are

the main source of energy for promoting respiration orobtaining metabolic energy and cellular biosynthesis(Stark amp Love 983090983088983088983091)

At harvest potato tubers are dormant and will notsprout even when placed in a favorable environmentdue to internal conditions such as the hormonalbalance of promoters and inhibitors As storage timegoes on the amount of promoters increases andsprout growth begins (Suttle 983090983088983088983092) In this contexttubers have three development stages apicaldominance multiple sprouting and senescence Inthe first dormancy is broken and a single sproutdevelops at the apex of the tuber in the secondseveral sprouts distributed throughout the tuberstart their growth and the last is characterized bylong branched thin and weak sprouts also turgorloss occurs in the tuber (Wiersema 983089983097983096983093)

The amount of sucrose in potato tubers is associated with the croprsquos degree of chemical maturity and thetype of processing to be performed after storage whilethe concentration of reducing sugars (glucose andfructose) is associated with frying quality (Sabba et al983090983088983088983095) Sugar content in tubers can be affected by the

variety its growth stage and environmental factors(crop management) (Thompson Love SowokinosThornton amp Shock 983090983088983088983096)

At harvest Van Es and Hartmans (983089983097983096983089) reportedconcentrations of 983089983090983096 total sugars and 983088983093983093 reducing sugars in potato tubers In other studiesRodriacuteguez-Saona and Wrolstad (983089983097983097983095) reported valuesof 983088983089983091 sucrose 983088983088983092 glucose and 983088983088983091 fructose

while Sabba et al (983090983088983088983095) indicate values of 983088983089983089 sucrose and 983088983088983093 glucose in dormant tubers

Introduccioacuten

La papa (Solanum tuberosum L) es el cuarto cultivo maacutesimportante en el mundo (Sonnewald amp Sonnewald983090983088983089983092) La formacioacuten de tubeacuterculos depende entreotros factores de la disponibilidad de asimilados yla habilidad de los tubeacuterculos para acumularlos Alinicio del desarrollo del cultivo los fotoasimilados se

destinan principalmente al desarrollo de hojas tallos y raiacuteces Sin embargo este comportamiento cambia aliniciar la tuberizacioacuten y crecimiento de los tubeacuterculosdebido a que incrementan la demanda de asimilados(Wolf 983089983097983097983091) de tal forma que los carbohidratosproducidos en las hojas a lo largo del ciclo del cultivoson traslocados a estas estructuras

Durante el inicio de la tuberizacioacuten el contenido dealmidoacuten es bajo y el de azuacutecares es alto ya que la tasade transporte de azuacutecares producidos en las hojassupera la de conversioacuten a almidoacuten de tal forma quecuando el tubeacuterculo alcanza la madurez fisioloacutegica

se obtiene la maacutexima acumulacioacuten de materia seca ymiacutenima cantidad de azuacutecares Eacutestos uacuteltimos constituyenla principal fuente de energiacutea para la respiracioacuten uobtencioacuten de energiacutea metaboacutelica y biosiacutentesis celular(Stark amp Love 983090983088983088983091)

Al momento de la cosecha los tubeacuterculos de papase encuentran en condiciones de reposo y no tienencapacidad de brotacioacuten aun cuando se coloquen enun ambiente favorable debido a condiciones internascomo el balance hormonal de promotores e inhibidoresConforme transcurre el almacenamiento la cantidadde promotores incrementa y comienza el crecimientodel brote (Suttle 983090983088983088983092) esto pretende definir tresestados de desarrollo del tubeacuterculo dominancia apicalbrotacioacuten muacuteltiple y senectud En la primera se rompeel reposo y se desarrolla un soacutelo brote en el aacutepice deltubeacuterculo en la segunda varios brotes distribuidosen el tubeacuterculo inician su crecimiento y la uacuteltima secaracteriza por largos brotes ramificados delgados ydeacutebiles ademaacutes ocurre la peacuterdida de turgencia en eltubeacuterculo (Wiersema 983089983097983096983093)

La cantidad de sacarosa en los tubeacuterculos de papase asocia al grado de madurez quiacutemica del cultivo

y al tipo de procesamiento a realizar despueacutes delalmacenamiento en tanto que la concentracioacuten deazuacutecares reductores (glucosa y fructosa) estaacute asociadacon la calidad en el freiacutedo (Sabba et al 983090983088983088983095) Elcontenido de azuacutecares en los tubeacuterculos puede serafectado por la variedad por su estado fenoloacutegico y porfactores ambientales (manejo del cultivo) (ThompsonLove Sowokinos Thornton amp Shock 983090983088983088983096)

Al momento de la cosecha de los tubeacuterculos de papa Van Es y Hartmans (983089983097983096983089) reportaron concentraciones

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

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The results of studies on the role of sugars intuber sprouting are ambiguous Dogras Siomosand Psomakelis (983089983097983096983097) observed an increase in theconcentration of total and reducing sugars in sproutedtubers of the variety Spunta whereas in cv Sebagothe content remained constant Debast et al (983090983088983089983089)indicate that the transport of sucrose to the tuberbuds is a prerequisite to induce sprouting and during

this stage the soluble sugar content decreases untilthe tuber sprouts reach approximately 983089 g dry matter(Viola et al 983090983088983088983095)

Therefore the objective of this research was to assessthe growth and yield of four potato varieties and todetermine the sugar content in tubers at differentphysiological ages and the effect these sugars have onseed-tuber sprouting

Materials and methods

This research was conducted in the municipality of San

Lorenzo Cuapiaxtla Tlaxcala (983089983097ordm 983089983096rsquo N 983097983095ordm 983092983094rsquo W and983090983092983089983088 masl) under irrigated conditions The regionrsquosclimate classification is C (Wo) (w) b (irsquo) C (Wo) (Garciacutea983089983097983096983096) and it has regosol-type soil

Four varieties were evaluated Alpha Atlantic Gigantand Vivaldi At planting minitubers of 983089983094 to 983090983096 mmin diameter in a state of apical dominance were usedThe experimental unit consisted of a row of 983093 m longand 983088983097983088 m wide with 983088983090983093 m spacing between plantsin each row 983090983088 minitubers were deposited at a depthof 983089983088 cm

At planting and 983090983088 days after emergence theminitubers were fertilized with the formula 983090983093983088-983094983088-983091983093983088-983097983089-983094983091 (N P K Ca and Mg) During the growingseason it was spray irrigated according to crop waterrequirements From emergence to tuber physiologicalmaturity the maximum and minimum air temperature(degC) was recorded with a Taylorreg model 983093983092983093983096 mercurythermometer using these data the mean temperature

was obtained

A randomized complete block experimental design with four replications was used and the experimental

unit consisted of four rows

From emergence the number of growing degree-days(GDD) accumulated at the beginning of each phaseand during each phenological stage of the crop wasdetermined with the classical residual method and983094 degC base temperature (Cao amp Tibbitts 983089983097983097983093) Theemergence phase (E) occurred when 983093983088 of the plantsof each experimental unit had emerged The stoloninitiation (SI) phase began when the stolon appearedon the main stem Tuber initiation or growth (T) was

de 983089983090983096 de azuacutecares totales y 983088983093983093 de azuacutecaresreductores En otros estudios Rodriacuteguez-Saona y

Wrolstad (983089983097983097983095) reportaron valores de 983088983089983091 desacarosa 983088983088983092 de glucosa y 983088983088983091 de fructosamientras que Sabba et al (983090983088983088983095) indican valores de983088983089983089 de sacarosa y 983088983088983093 de glucosa en tubeacuterculos enestado de reposo

Los resultados de estudios sobre el papel de losazuacutecares en la brotacioacuten de los tubeacuterculos sonpoco claros Dogras Siomos y Psomakelis (983089983097983096983097)observaron incremento en la concentracioacuten deazuacutecares totales y reductores en tubeacuterculos brotadosde la variedad Spunta en tanto que en cv Sebagoel contenido se mantuvo constante Debast et al(983090983088983089983089) indican que el transporte de sacarosa a las

yemas del tubeacuterculo es prerrequisito para inducir labrotacioacuten y durante eacutesta el contenido de azuacutecaressolubles disminuye hasta que los brotes del tubeacuterculoalcanzan aproximadamente 983089 g de materia seca(Viola et al 983090983088983088983095)

Por lo anterior la presente investigacioacuten tuvo comoobjetivo evaluar el crecimiento y rendimiento de cuatro

variedades de papa determinar el contenido de azuacutecaresen tubeacuterculos con diferente edad fisioloacutegica y el efectoque tienen estos en la brotacioacuten de la semilla-tubeacuterculo

Materiales y meacutetodos

La presente investigacioacuten se realizoacute en el municipiode San Lorenzo Cuapiaxtla Tlaxcala (983089983097ordm 983089983096rsquo N 983097983095ordm 983092983094rsquoO y 983090983092983089983088 msnm) bajo condiciones de riego La regioacutenposee clima C (Wo) (w) b (irsquo) C (Wo) (Garciacutea 983089983097983096983096) ysuelo de tipo regosol

Se evaluaron las variedades Alpha Atlantic Gigant y Vivaldi En la siembra se usaron minitubeacuterculos de 983089983094a 983090983096 mm de diaacutemetro en estado de dominancia apicalla unidad experimental consistioacute en un surco de 983093 m delongitud con 983088983097983088 m de ancho y 983088983090983093 m entre plantasen cada surco se depositaron 983090983088 minitubeacuterculos a 983089983088cm de profundidad

En la siembra y 983090983088 diacuteas despueacutes de la emergencia sefertilizoacute con la foacutermula 983090983093983088-983094983088-983091983093983088-983097983089-983094983091 (N P K Ca y

Mg) Durante el ciclo de cultivo se irrigoacute por aspersioacutende acuerdo con las necesidades hiacutedricas del mismoDesde la emergencia y hasta la madurez fisioloacutegicadel tubeacuterculo se registroacute la temperatura maacutexima ymiacutenima del aire (ordmC) con un termoacutemetro de columna demercurio marca Taylorreg modelo 983093983092983093983096 con esos datos seobtuvo la temperatura media

El disentildeo experimental fue bloques completos al azarcon cuatro repeticiones y la unidad experimentalestuvo constituida por cuatro surcos

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983091983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

identified as the time at which the apex of the stolonthickened Tuber physiological maturity (TM) was

visually identified by the loss of the foliagersquos greencolor or onset of senescence

The vegetative stage (VS) was considered as the timefrom emergence to the beginning of the stolonphase during which shoot growth and root system

establishment occur The initial reproductive stage (RS983089) includes the interval between the onset of the stolonstage and tuberization and is characterized by stolongeneration and growth The final reproductive stage(RS 983090) runs from the beginning of tuberization to tuberphysiological maturity where tuber growth occurred(Morales-Fernaacutendez et al 983090983088983089983089)

At harvest the number (NCTP) and weight (WCTP g) ofcommercial tubers per plant (with a greater than 983090983088-mmdiameter) were quantified as were the number (NNCTP)and weight (WNCTP g) of non-commercial tubers perplant (with a 983090983088-mm diameter) characterized by being

deformed cracked and green The total number of tubersper plant (TNTP) was obtained by adding NCTP and NNCTPThe total weight of tubers per plant (TWTP g) was the sumof WCTP plus WNCTP The average tuber weight per plant(ATWP) was obtained by dividing TWTP by TNTP

Storage conditions

At harvest the tubers of four randomly-selected plantsper experimental unit were placed in paper bags to belater stored in the laboratory with ambient light underthe temperature and relative humidity conditions shownin Table 983089 until showing maximum sprout developmentIt was considered that the tubers contained in thepaper bags left dormancy when in 983093983088 of them the mainsprout measured 983092 plusmn 983090 mm Sprout length (mm) anddry weight (mg) were recorded in five tubers selected

A partir de la emergencia se determinoacute el nuacutemerode grados diacutea (GD) acumulados al inicio de cada fase

y durante cada etapa fenoloacutegica del cultivo con elmeacutetodo residual claacutesico y la temperatura base de 983094 ordmC (Caoamp Tibbitts 983089983097983097983093) La fase de emergencia (E) ocurrioacute cuando983093983088 de las plantas de cada unidad experimental habiacuteanemergido El inicio de estoloneo (ES) fue cuando aparecioacuteel estoloacuten en el tallo principal El inicio de tuberizacioacuten

o de crecimiento del tubeacuterculo (T) se identificoacute como elmomento en que engrosoacute el aacutepice del estoloacuten La madurezfisioloacutegica del tubeacuterculo (MT) se identificoacute visualmentepor la peacuterdida del color verde del follaje o inicio de lasenescencia de eacuteste

La etapa vegetativa (EV) se consideroacute como el periodotranscurrido desde la emergencia hasta el inicio deestoloneo durante el cual ocurre el crecimiento de laparte aeacuterea y el establecimiento del sistema radicalLa etapa reproductiva inicial (ER 983089) incluyoacute el intervaloentre el inicio de estoloneo y el de tuberizacioacuten y secaracterizoacute por la generacioacuten y crecimiento de estolones

La etapa reproductiva final (ER 983090) va desde el inicio de latuberizacioacuten hasta la madurez fisioloacutegica del tubeacuterculodonde ocurrioacute el crecimiento de este oacutergano (Morales-Fernaacutendez et al 983090983088983089983089)

Durante la cosecha se cuantificoacute el nuacutemero detubeacuterculos por planta (NTCP) y peso de tubeacuterculos porplanta (PTCP g) de tubeacuterculos comerciales por planta(con diaacutemetro mayor de 983090983088 mm) nuacutemero (NTNCP)

y peso (PTNCP g) de tubeacuterculos no comerciales porplanta (con diaacutemetro menor de 983090983088 mm) deformesrajados y verdes El nuacutemero total de tubeacuterculos porplanta (NTTP) que se obtuvo al sumar NTCP maacutesNTNCP El peso total de tubeacuterculos por planta (PTTPg) fue la suma de PTCP maacutes PTNCP El peso promediode tubeacuterculo por planta (PPTP) se obtuvo al dividirPTTP entre NTTP

Table 983089 Air temperature and relative humidity recorded during storage of potato seed-tubers after harvest

Cuadro 983089 Temperatura del aire y humedad relativa registradas durante el almacenamiento de la semilla-

tubeacuterculo de papa despueacutes de la cosecha

ELEMENT frasl ELEMENTO

983090983088983088983096 983090983088983088983097

AGO frasl

AUGSEP OCT NOV

DIC frasl

DECENE FEB MAR

ABR frasl

APR MAY JUN

Temperature (ordmC) frasl Temperatura (ordmC)

Maximum frasl Maacutexima 983090983094 983090983092 983090983089 983089983095 983089983094 983089983093 983089983095 983090983089 983090983092 983090983094 983090983095

Minimum frasl Miacutenima 983089983096 983089983096 983089983094 983089983090 983089983090 983089983089 983089983092 983089983092 983089983095 983090983089 983090983088

Mean frasl Media 983090983090 983090983089 983089983096 983089983092 983089983092 983089983091 983089983093 983089983095 983090983088 983090983091 983090983091

Humedad relativa () frasl

Relative humidity ()

Maximum frasl Maacutexima 983092983090 983092983092 983091983093 983091983088 983091983088 983091983089 983091983091 983091983090 983091983092 983091983094 983092983094

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at random from the experimental unit during apicaldominance (a single sprout) multiple sprouting (morethan one sprout) and senescence (more than onesprout and turgor loss in tubers) On each occasion the longest sprout was measured and from multiplesprouting all sprouts were measured for dry weight

Quantification of soluble carbohydrates by high-

performance liquid chromatography (HPLC)

Sample collection

Four samplings were performed during the storageperiod the first during dormancy and the other threeduring the physiological ages of apical dominancemultiple sprouting and senescence of the tuber ofeach variety On each occasion a tuber contained in thepaper bags was taken and a sample obtained from it

with two replications being made for the larger tubers Average tuber weights for the sampling process rangedbetween 983089983093983090 and 983089983096983089 g (Table 983090)

From the central part of each tuber between the basaland apical end a sample of 983093 plusmn 983089 g was extracted andplaced in liquid nitrogen (-983089983097983094 degC) until analysis

Extraction of soluble carbohydrates

The frozen tuber samples were individually ground ina blender with 983090983088 mL of 983095983088 HPLC-grade ethanol and

Condiciones de almacenamiento

En la cosecha los tubeacuterculos de cuatro plantas por unidadexperimental seleccionadas al azar fueron colocados enbolsas de papel para ser almacenados en laboratorio conluz ambiental bajo las condiciones de temperatura yhumedad relativa que se muestran en el Cuadro 983089 hastaque mostraron el maacuteximo desarrollo de brote

Se consideroacute que los tubeacuterculos contenidos en lasbolsas de papel dejaron la condicioacuten de reposo cuandoen 983093983088 de estos el brote principal midioacute 983092 plusmn 983090 mm Lalongitud (mm) y peso seco (mg) del brote se registroacuteen cinco tubeacuterculos elegidos al azar de la unidadexperimental en dominancia apical (un solo brote)brotacioacuten muacuteltiple (maacutes de un brote) y senescencia (maacutesde un brote y peacuterdida de turgencia de los tubeacuterculos)En cada ocasioacuten se midioacute el brote de mayor longitud ya partir de la brotacioacuten muacuteltiple se consideraron todoslos brotes para medir el peso seco

Cuantificacioacuten de carbohidratos solubles por croma-tografiacutea liacutequida de alta resolucioacuten (HPLC)

Obtencioacuten de la muestra

Durante el periodo de almacenamiento se realizaroncuatro muestreos el primero en condiciones dereposo y los otros tres correspondieron a las edadesfisioloacutegicas de dominancia apical brotacioacuten muacuteltiple

Table 983090 Seed-tuber weight at different physiological states of four potato varieties

Cuadro 983090 Peso de semilla-tubeacuterculo en diferentes estados fisioloacutegicos de cuatro variedades de papa

Seed-tuber hysiological state frasl Estado fisioloacutegico de semilla-tubeacuterculo

Variety frasl

VariedadDormancy frasl Reposo

Apical dominance frasl

Dominancia apical

Multiple sprouting frasl

Brotacioacuten muacuteltiple

Senescence frasl

Senescencia

DAHz frasl

DDCz

Weight y (g) frasl

Peso y (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

Alpha 983090983090 983089983097983089 983089983089983090 983089983096983090 983089983092983096 983089983094983091 983090983090983091 983089983093983088

Atlantic 983090983090 983089983092983090 983089983092983095 983089983093983089 983090983089983094 983089983092983093 983090983096983088 983089983091983088

Gigant 983090983090 983090983088983093 983095983095 983090983091983096 983097983096 983090983090983096 983090983090983091 983090983088983088

Vivaldi 983090983090 983089983096983096 983097983096 983089983092983094 983089983092983096 983089983092983097 983090983090983091 983089983091983088

Mean fraslMedia

983090983090 983089983096983089 983089983088983097 983089983095983097 983089983093983091 983089983095983089 983090983091983095 983089983093983090

zDDC days after harvest y average value of two tubers

zDDC diacuteas despueacutes de la cosecha y valor promedio de dos tubeacuterculos

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983089983091983092 Growth yield and sugar

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brought to boiling temperature for 983093 min The extract was decanted and a second extraction was made fromthe residue with 983089983088 mL of ethanol at the end theextracts were mixed Subsequently the extract wascentrifuged for 983089983093 min at 983092983088983088983088 x g in a HERLE modelZ983090983091983088A centrifuge (Labnet International Inc Edison NJ

USA) filtered with Whatman No 983092 paper and mannitol was added at a concentration of 983092 mgmiddotmL-983089 as internal

standard Finally the extract was gauged to 983090983093 mL withthe same kind of solvent used for the extraction

Then 983089983088 mL of the extract were taken and concentratedto dryness with reduced pressure and low temperature(983092983088 degC) in a BUCHI R-983090983089983093 rotary evaporator (Switzerland)The residue was resuspended in 983090 mL of HPLC-grade

water in order to pass it through a column containingion-exchange resins one basic (983088983093 mL of Dowex - 983089 x983096 - Fluka) and the other acidic (983088983093 mL of Dowex - 983093983088W

X983096 - Fluka) at a ratio of 983089983089 (ww) The purified sample was passed through a 983088983092983093-μm nylon acrodisc in orderto collect the sample in a 983089983093 mL vial

HPLC sugar analysis

A Perkin Elmer series 983090983088983088 high-performance liquidchromatography system (Boston Ma USA) consistingof an autosampler quaternary pump degasserrefractive index detector and a column oven wasused The system is operated with TotalChrom version983094983090983089 software A Rezex RCM-monosaccharide Ca+983090 (983096 )column 983091983088 cm in length and 983095983096 mm in diameter wasused As the mobile phase HPLC-grade water was usedThe flow rate was 983088983094 mLmiddotmin-983089 the sample injection

volume 983090983088 μL and the running time 983090983093 min Columntemperature was maintained at 983096983093 degC Pattern curves

were prepared using commercial sucrose glucose andfructose standards (Sigma MN) at concentrations of983088983093 983089983088 983090983093 and 983093 mgmiddotmL-983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) At each point of the curves it was injected at leastthree times to obtain the data of the area associated

with each concentration area using this informationthe regression equations for each sugar were obtained

Analyses of variance and regression were performed with the data obtained Also Tukeyrsquos test was appliedusing the Statistical Analysis System package (SAS 983090983088983088983092)

Results and discussion

Temperature

The air temperature fluctuated during the growingseason (Figure 983089) with the maximum varying between983089983095 and 983091983094 degC Temperatures above 983090983091 degC were recordedfrom 983096 to 983093983092 days after emergence and coincided

with the vegetative (VS) and initial reproductive (RS983089)stages and half of the final reproductive stage (RS 983090)

y senescencia del tubeacuterculo de cada variedad En cadaocasioacuten se tomoacute un tubeacuterculo contenido en las bolsasde papel el de mayor tamantildeo en dos repeticiones conpesos promedio entre 983089983093983090 y 983089983096983089 g (Cuadro 983090) del que seobtuvo la muestra

De la parte central de cada tubeacuterculo entre el extremobasal y apical se extrajo una muestra de 983093 plusmn 983089 g que se

colocoacute en nitroacutegeno liacutequido (-983089983097983094 ordmC) hasta su anaacutelisis

Extraccioacuten de carbohidratos solubles

Las muestras congeladas de tubeacuterculo fueron molidasen licuadora individualmente con 983090983088 mL de etanolgrado HPLC a 983095983088 y llevadas a temperatura deebullicioacuten durante 983093 min El extracto se decantoacute yal residuo se le realizoacute una segunda extraccioacuten con983089983088 mL de etanol al final los extractos se mezclaronPosteriormente el extracto se centrifugoacute por 983089983093 mina 983092983088983088983088 x g en una centriacutefuga HERLE modelo Z983090983091983088A(Labnet International Inc Edison NJ USA) se filtroacute

con papel Whatman nuacutemero 983092 y se agregoacute manitol enconcentracioacuten de 983092 mgmiddotmL-983089 como estaacutendar internoFinalmente el extracto se aforoacute a 983090983093 mL con el mismotipo de solvente empleado para la extraccioacuten

Del extracto aforado se tomaron 983089983088 mL paraconcentrar a sequedad con presioacuten reducida y bajatemperatura (983092983088 ordmC) en un rotoevaporador BUCHIR-983090983089983093 (Switzerland) El residuo se resuspendioacute en 983090 mLde agua grado HPLC para pasarlo por una columnaque conteniacutea resinas intercambiadoras de iones unabaacutesica (983088983093 mL de Dowex - 983089 x 983096 - Fluka) y la otra aacutecida(983088983093 mL de Dowex - 983093983088W X983096 - Fluka) en proporcioacuten 983089983089(pp) La muestra purificada se pasoacute por un acrodiscode nylon de 983088983092983093 μm con el fin de colectar la muestraen un vial de 983089983093 mL

Anaacutelisis de azuacutecares en HPLC

Se utilizoacute un equipo de cromatografiacutea liacutequida de altaresolucioacuten serie 983090983088983088 Perkin Elmer (Boston Ma USA)compuesto por un auto muestreador bomba cuaternariadegasificador detector de iacutendice de refraccioacuten y hornopara columnas El sistema se operoacute con el programacomputacional TotalChrom versioacuten 983094983090983089 Se usoacute una

columna Rezex RCM-monosaccharide Ca+983090

(983096 ) de 983091983088cm de longitud y 983095983096 mm de diaacutemetro Como fase moacutevilse empleoacute agua grado HPLC La velocidad de flujo fuede 983088983094 mLmiddotmin-983089 el volumen de muestra inyectado de983090983088 μL y el tiempo de corrida 983090983093 min La temperatura dela columna se mantuvo a 983096983093 ordmC Se elaboraron curvaspatroacuten usando estaacutendares comerciales de sacarosaglucosa y fructosa (Sigma MN) en concentraciones de983088983093 983089983088 983090983093 y 983093 mgmiddotmL -983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) En cada punto de las curvas se inyectoacute al menostres veces para obtener los datos del aacuterea asociada a

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The minimum ranged between 983095 and 983089983094 degC and eventhough in some periods it was less than 983089983088 degC it did notaffect plant growth and yield since it is considered thatpotato tolerates low temperatures with 983094 degC being itsbase temperature (Cao amp Tibbitts 983089983097983097983093)

The mean temperature was 983089983097 degC similar to thatreported by Cao and Tibbitts (983089983097983097983092) to obtain high

biomass and tuber production (983090983088 degC) Struik Haverkort Vreugdengil Bus and Dankert (983089983097983097983088) stress theimportance of high temperatures during RS 983089 as theypromote stolon branching an important condition for

yield because the number of tubers formed is closelyrelated to the number of stolons (Haverkort Van De

Waart amp Bodlaender 983089983097983097983088)

Crop Phenology

In general there were differential genotypic effects ( P le983088983088983093) in crop phenology The variety Alpha required moreGDD to reach the stolon (983089983094983095) tuberization (983090983094983089) and tuber

physiological maturity (983089983093983092) stages of the tuber its lifecycle was significantly longer than in the other varieties

Vivaldi was the variety that reached maturity the fastestbecause it needed 983089983093 fewer GDD than Alpha (Table 983091)

cada concentracioacuten con esta informacioacuten se obtuvieronlas ecuaciones de regresioacuten para cada azuacutecar

Con los datos obtenidos se realizaron un anaacutelisis de varianza y de regresioacuten ademaacutes se aplicaron pruebas decomparacioacuten de medias de Tukey para ello se empleoacuteel paquete Statistical Analysis System (SAS 983090983088983088983092)

Resultados y discusioacuten

Temperatura

La temperatura del aire fue fluctuante durante el ciclode cultivo (Figura 983089) la maacutexima varioacute entre 983089983095 y 983091983094 ordmCSuperiores a 983090983091 ordmC se registraron desde los 983096 hasta los983093983092 diacuteas despueacutes de la emergencia y coincidieron con laetapa vegetativa (EV) reproductiva inicial (ER 983089) y mitadde la reproductiva final (ER 983090) La miacutenima osciloacute entre 983095

y 983089983094 ordmC y a pesar de que en algunos periodos fue inferiora 983089983088 ordmC no afectoacute al crecimiento y rendimiento de laplanta ya que se considera que la papa resiste bajas

temperaturas siendo su base 983094 ordmC (Cao amp Tibbitts 983089983097983097983093)

La temperatura media fue de 983089983097 ordmC similar a lareportada por Cao y Tibbitts (983089983097983097983092) para obtener alta

Figure 983089 Air temperature recorded during the potato growing season under field conditions San Lorenzo

Cuapiaxtla Tlax Spring-Summer Cycle 983090983088983088983096 The arrows indicate the phases of emergence (E) stolon

initiation (SI) tuberization (T) and tuber physiological maturity (TM) and the intervals between them

at the vegetative (VS) initial reproductive (RS 983089) and final reproductive (RS 983090) stages Average values of

four varieties

Figura 983089 Temperatura del aire registrada durante el ciclo de cultivo de papa en condiciones de campo San

Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096 Las flechas indican las fases de emergencia

(E) inicio de estoloneo (ES) tuberizacioacuten (T) y madurez fisioloacutegica del tubeacuterculo (MT) y los intervalos

entre estas a las etapas vegetativa (EV) reproductiva inicial (ER 983089) y reproductiva final (ER 983090) Valores

promedio de cuatro variedades

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The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

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Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

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durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

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Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

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significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

Ap Rees T amp Morrell S (983089983097983097983088) Carbohydrate metabolismin developing potatoes American Journal of Potato

Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

change patterns affect growth and tuberization inpotatoes Journal of the American Society for Horticultural

Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 218

983089983091983088 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Introduction

The potato (Solanum tuberosum L) is the fourth mostimportant crop in the world (Sonnewald amp Sonnewald983090983088983089983092) Tuber formation depends on among other factorsthe availability of assimilates and the ability of tubersto accumulate them At the start of crop developmentphotoassimilates are mainly directed towards the

development of leaves stems and roots However thisbehavior changes with tuber initiation and growthbecause they increase the demand for assimilates (Wolf983089983097983097983091) and consequently the carbohydrates producedin the leaves throughout the crop cycle are translocatedto tuber structures

During tuber initiation starch content is low but thatof sugars is high since the rate of transport of sugarsproduced in the leaves exceeds the rate of conversion tostarch As a result when the tuber reaches physiologicalmaturity the maximum dry matter accumulation andminimum amount of sugars are obtained The latter are

the main source of energy for promoting respiration orobtaining metabolic energy and cellular biosynthesis(Stark amp Love 983090983088983088983091)

At harvest potato tubers are dormant and will notsprout even when placed in a favorable environmentdue to internal conditions such as the hormonalbalance of promoters and inhibitors As storage timegoes on the amount of promoters increases andsprout growth begins (Suttle 983090983088983088983092) In this contexttubers have three development stages apicaldominance multiple sprouting and senescence Inthe first dormancy is broken and a single sproutdevelops at the apex of the tuber in the secondseveral sprouts distributed throughout the tuberstart their growth and the last is characterized bylong branched thin and weak sprouts also turgorloss occurs in the tuber (Wiersema 983089983097983096983093)

The amount of sucrose in potato tubers is associated with the croprsquos degree of chemical maturity and thetype of processing to be performed after storage whilethe concentration of reducing sugars (glucose andfructose) is associated with frying quality (Sabba et al983090983088983088983095) Sugar content in tubers can be affected by the

variety its growth stage and environmental factors(crop management) (Thompson Love SowokinosThornton amp Shock 983090983088983088983096)

At harvest Van Es and Hartmans (983089983097983096983089) reportedconcentrations of 983089983090983096 total sugars and 983088983093983093 reducing sugars in potato tubers In other studiesRodriacuteguez-Saona and Wrolstad (983089983097983097983095) reported valuesof 983088983089983091 sucrose 983088983088983092 glucose and 983088983088983091 fructose

while Sabba et al (983090983088983088983095) indicate values of 983088983089983089 sucrose and 983088983088983093 glucose in dormant tubers

Introduccioacuten

La papa (Solanum tuberosum L) es el cuarto cultivo maacutesimportante en el mundo (Sonnewald amp Sonnewald983090983088983089983092) La formacioacuten de tubeacuterculos depende entreotros factores de la disponibilidad de asimilados yla habilidad de los tubeacuterculos para acumularlos Alinicio del desarrollo del cultivo los fotoasimilados se

destinan principalmente al desarrollo de hojas tallos y raiacuteces Sin embargo este comportamiento cambia aliniciar la tuberizacioacuten y crecimiento de los tubeacuterculosdebido a que incrementan la demanda de asimilados(Wolf 983089983097983097983091) de tal forma que los carbohidratosproducidos en las hojas a lo largo del ciclo del cultivoson traslocados a estas estructuras

Durante el inicio de la tuberizacioacuten el contenido dealmidoacuten es bajo y el de azuacutecares es alto ya que la tasade transporte de azuacutecares producidos en las hojassupera la de conversioacuten a almidoacuten de tal forma quecuando el tubeacuterculo alcanza la madurez fisioloacutegica

se obtiene la maacutexima acumulacioacuten de materia seca ymiacutenima cantidad de azuacutecares Eacutestos uacuteltimos constituyenla principal fuente de energiacutea para la respiracioacuten uobtencioacuten de energiacutea metaboacutelica y biosiacutentesis celular(Stark amp Love 983090983088983088983091)

Al momento de la cosecha los tubeacuterculos de papase encuentran en condiciones de reposo y no tienencapacidad de brotacioacuten aun cuando se coloquen enun ambiente favorable debido a condiciones internascomo el balance hormonal de promotores e inhibidoresConforme transcurre el almacenamiento la cantidadde promotores incrementa y comienza el crecimientodel brote (Suttle 983090983088983088983092) esto pretende definir tresestados de desarrollo del tubeacuterculo dominancia apicalbrotacioacuten muacuteltiple y senectud En la primera se rompeel reposo y se desarrolla un soacutelo brote en el aacutepice deltubeacuterculo en la segunda varios brotes distribuidosen el tubeacuterculo inician su crecimiento y la uacuteltima secaracteriza por largos brotes ramificados delgados ydeacutebiles ademaacutes ocurre la peacuterdida de turgencia en eltubeacuterculo (Wiersema 983089983097983096983093)

La cantidad de sacarosa en los tubeacuterculos de papase asocia al grado de madurez quiacutemica del cultivo

y al tipo de procesamiento a realizar despueacutes delalmacenamiento en tanto que la concentracioacuten deazuacutecares reductores (glucosa y fructosa) estaacute asociadacon la calidad en el freiacutedo (Sabba et al 983090983088983088983095) Elcontenido de azuacutecares en los tubeacuterculos puede serafectado por la variedad por su estado fenoloacutegico y porfactores ambientales (manejo del cultivo) (ThompsonLove Sowokinos Thornton amp Shock 983090983088983088983096)

Al momento de la cosecha de los tubeacuterculos de papa Van Es y Hartmans (983089983097983096983089) reportaron concentraciones

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 318

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

The results of studies on the role of sugars intuber sprouting are ambiguous Dogras Siomosand Psomakelis (983089983097983096983097) observed an increase in theconcentration of total and reducing sugars in sproutedtubers of the variety Spunta whereas in cv Sebagothe content remained constant Debast et al (983090983088983089983089)indicate that the transport of sucrose to the tuberbuds is a prerequisite to induce sprouting and during

this stage the soluble sugar content decreases untilthe tuber sprouts reach approximately 983089 g dry matter(Viola et al 983090983088983088983095)

Therefore the objective of this research was to assessthe growth and yield of four potato varieties and todetermine the sugar content in tubers at differentphysiological ages and the effect these sugars have onseed-tuber sprouting

Materials and methods

This research was conducted in the municipality of San

Lorenzo Cuapiaxtla Tlaxcala (983089983097ordm 983089983096rsquo N 983097983095ordm 983092983094rsquo W and983090983092983089983088 masl) under irrigated conditions The regionrsquosclimate classification is C (Wo) (w) b (irsquo) C (Wo) (Garciacutea983089983097983096983096) and it has regosol-type soil

Four varieties were evaluated Alpha Atlantic Gigantand Vivaldi At planting minitubers of 983089983094 to 983090983096 mmin diameter in a state of apical dominance were usedThe experimental unit consisted of a row of 983093 m longand 983088983097983088 m wide with 983088983090983093 m spacing between plantsin each row 983090983088 minitubers were deposited at a depthof 983089983088 cm

At planting and 983090983088 days after emergence theminitubers were fertilized with the formula 983090983093983088-983094983088-983091983093983088-983097983089-983094983091 (N P K Ca and Mg) During the growingseason it was spray irrigated according to crop waterrequirements From emergence to tuber physiologicalmaturity the maximum and minimum air temperature(degC) was recorded with a Taylorreg model 983093983092983093983096 mercurythermometer using these data the mean temperature

was obtained

A randomized complete block experimental design with four replications was used and the experimental

unit consisted of four rows

From emergence the number of growing degree-days(GDD) accumulated at the beginning of each phaseand during each phenological stage of the crop wasdetermined with the classical residual method and983094 degC base temperature (Cao amp Tibbitts 983089983097983097983093) Theemergence phase (E) occurred when 983093983088 of the plantsof each experimental unit had emerged The stoloninitiation (SI) phase began when the stolon appearedon the main stem Tuber initiation or growth (T) was

de 983089983090983096 de azuacutecares totales y 983088983093983093 de azuacutecaresreductores En otros estudios Rodriacuteguez-Saona y

Wrolstad (983089983097983097983095) reportaron valores de 983088983089983091 desacarosa 983088983088983092 de glucosa y 983088983088983091 de fructosamientras que Sabba et al (983090983088983088983095) indican valores de983088983089983089 de sacarosa y 983088983088983093 de glucosa en tubeacuterculos enestado de reposo

Los resultados de estudios sobre el papel de losazuacutecares en la brotacioacuten de los tubeacuterculos sonpoco claros Dogras Siomos y Psomakelis (983089983097983096983097)observaron incremento en la concentracioacuten deazuacutecares totales y reductores en tubeacuterculos brotadosde la variedad Spunta en tanto que en cv Sebagoel contenido se mantuvo constante Debast et al(983090983088983089983089) indican que el transporte de sacarosa a las

yemas del tubeacuterculo es prerrequisito para inducir labrotacioacuten y durante eacutesta el contenido de azuacutecaressolubles disminuye hasta que los brotes del tubeacuterculoalcanzan aproximadamente 983089 g de materia seca(Viola et al 983090983088983088983095)

Por lo anterior la presente investigacioacuten tuvo comoobjetivo evaluar el crecimiento y rendimiento de cuatro

variedades de papa determinar el contenido de azuacutecaresen tubeacuterculos con diferente edad fisioloacutegica y el efectoque tienen estos en la brotacioacuten de la semilla-tubeacuterculo

Materiales y meacutetodos

La presente investigacioacuten se realizoacute en el municipiode San Lorenzo Cuapiaxtla Tlaxcala (983089983097ordm 983089983096rsquo N 983097983095ordm 983092983094rsquoO y 983090983092983089983088 msnm) bajo condiciones de riego La regioacutenposee clima C (Wo) (w) b (irsquo) C (Wo) (Garciacutea 983089983097983096983096) ysuelo de tipo regosol

Se evaluaron las variedades Alpha Atlantic Gigant y Vivaldi En la siembra se usaron minitubeacuterculos de 983089983094a 983090983096 mm de diaacutemetro en estado de dominancia apicalla unidad experimental consistioacute en un surco de 983093 m delongitud con 983088983097983088 m de ancho y 983088983090983093 m entre plantasen cada surco se depositaron 983090983088 minitubeacuterculos a 983089983088cm de profundidad

En la siembra y 983090983088 diacuteas despueacutes de la emergencia sefertilizoacute con la foacutermula 983090983093983088-983094983088-983091983093983088-983097983089-983094983091 (N P K Ca y

Mg) Durante el ciclo de cultivo se irrigoacute por aspersioacutende acuerdo con las necesidades hiacutedricas del mismoDesde la emergencia y hasta la madurez fisioloacutegicadel tubeacuterculo se registroacute la temperatura maacutexima ymiacutenima del aire (ordmC) con un termoacutemetro de columna demercurio marca Taylorreg modelo 983093983092983093983096 con esos datos seobtuvo la temperatura media

El disentildeo experimental fue bloques completos al azarcon cuatro repeticiones y la unidad experimentalestuvo constituida por cuatro surcos

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identified as the time at which the apex of the stolonthickened Tuber physiological maturity (TM) was

visually identified by the loss of the foliagersquos greencolor or onset of senescence

The vegetative stage (VS) was considered as the timefrom emergence to the beginning of the stolonphase during which shoot growth and root system

establishment occur The initial reproductive stage (RS983089) includes the interval between the onset of the stolonstage and tuberization and is characterized by stolongeneration and growth The final reproductive stage(RS 983090) runs from the beginning of tuberization to tuberphysiological maturity where tuber growth occurred(Morales-Fernaacutendez et al 983090983088983089983089)

At harvest the number (NCTP) and weight (WCTP g) ofcommercial tubers per plant (with a greater than 983090983088-mmdiameter) were quantified as were the number (NNCTP)and weight (WNCTP g) of non-commercial tubers perplant (with a 983090983088-mm diameter) characterized by being

deformed cracked and green The total number of tubersper plant (TNTP) was obtained by adding NCTP and NNCTPThe total weight of tubers per plant (TWTP g) was the sumof WCTP plus WNCTP The average tuber weight per plant(ATWP) was obtained by dividing TWTP by TNTP

Storage conditions

At harvest the tubers of four randomly-selected plantsper experimental unit were placed in paper bags to belater stored in the laboratory with ambient light underthe temperature and relative humidity conditions shownin Table 983089 until showing maximum sprout developmentIt was considered that the tubers contained in thepaper bags left dormancy when in 983093983088 of them the mainsprout measured 983092 plusmn 983090 mm Sprout length (mm) anddry weight (mg) were recorded in five tubers selected

A partir de la emergencia se determinoacute el nuacutemerode grados diacutea (GD) acumulados al inicio de cada fase

y durante cada etapa fenoloacutegica del cultivo con elmeacutetodo residual claacutesico y la temperatura base de 983094 ordmC (Caoamp Tibbitts 983089983097983097983093) La fase de emergencia (E) ocurrioacute cuando983093983088 de las plantas de cada unidad experimental habiacuteanemergido El inicio de estoloneo (ES) fue cuando aparecioacuteel estoloacuten en el tallo principal El inicio de tuberizacioacuten

o de crecimiento del tubeacuterculo (T) se identificoacute como elmomento en que engrosoacute el aacutepice del estoloacuten La madurezfisioloacutegica del tubeacuterculo (MT) se identificoacute visualmentepor la peacuterdida del color verde del follaje o inicio de lasenescencia de eacuteste

La etapa vegetativa (EV) se consideroacute como el periodotranscurrido desde la emergencia hasta el inicio deestoloneo durante el cual ocurre el crecimiento de laparte aeacuterea y el establecimiento del sistema radicalLa etapa reproductiva inicial (ER 983089) incluyoacute el intervaloentre el inicio de estoloneo y el de tuberizacioacuten y secaracterizoacute por la generacioacuten y crecimiento de estolones

La etapa reproductiva final (ER 983090) va desde el inicio de latuberizacioacuten hasta la madurez fisioloacutegica del tubeacuterculodonde ocurrioacute el crecimiento de este oacutergano (Morales-Fernaacutendez et al 983090983088983089983089)

Durante la cosecha se cuantificoacute el nuacutemero detubeacuterculos por planta (NTCP) y peso de tubeacuterculos porplanta (PTCP g) de tubeacuterculos comerciales por planta(con diaacutemetro mayor de 983090983088 mm) nuacutemero (NTNCP)

y peso (PTNCP g) de tubeacuterculos no comerciales porplanta (con diaacutemetro menor de 983090983088 mm) deformesrajados y verdes El nuacutemero total de tubeacuterculos porplanta (NTTP) que se obtuvo al sumar NTCP maacutesNTNCP El peso total de tubeacuterculos por planta (PTTPg) fue la suma de PTCP maacutes PTNCP El peso promediode tubeacuterculo por planta (PPTP) se obtuvo al dividirPTTP entre NTTP

Table 983089 Air temperature and relative humidity recorded during storage of potato seed-tubers after harvest

Cuadro 983089 Temperatura del aire y humedad relativa registradas durante el almacenamiento de la semilla-

tubeacuterculo de papa despueacutes de la cosecha

ELEMENT frasl ELEMENTO

983090983088983088983096 983090983088983088983097

AGO frasl

AUGSEP OCT NOV

DIC frasl

DECENE FEB MAR

ABR frasl

APR MAY JUN

Temperature (ordmC) frasl Temperatura (ordmC)

Maximum frasl Maacutexima 983090983094 983090983092 983090983089 983089983095 983089983094 983089983093 983089983095 983090983089 983090983092 983090983094 983090983095

Minimum frasl Miacutenima 983089983096 983089983096 983089983094 983089983090 983089983090 983089983089 983089983092 983089983092 983089983095 983090983089 983090983088

Mean frasl Media 983090983090 983090983089 983089983096 983089983092 983089983092 983089983091 983089983093 983089983095 983090983088 983090983091 983090983091

Humedad relativa () frasl

Relative humidity ()

Maximum frasl Maacutexima 983092983090 983092983092 983091983093 983091983088 983091983088 983091983089 983091983091 983091983090 983091983092 983091983094 983092983094

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at random from the experimental unit during apicaldominance (a single sprout) multiple sprouting (morethan one sprout) and senescence (more than onesprout and turgor loss in tubers) On each occasion the longest sprout was measured and from multiplesprouting all sprouts were measured for dry weight

Quantification of soluble carbohydrates by high-

performance liquid chromatography (HPLC)

Sample collection

Four samplings were performed during the storageperiod the first during dormancy and the other threeduring the physiological ages of apical dominancemultiple sprouting and senescence of the tuber ofeach variety On each occasion a tuber contained in thepaper bags was taken and a sample obtained from it

with two replications being made for the larger tubers Average tuber weights for the sampling process rangedbetween 983089983093983090 and 983089983096983089 g (Table 983090)

From the central part of each tuber between the basaland apical end a sample of 983093 plusmn 983089 g was extracted andplaced in liquid nitrogen (-983089983097983094 degC) until analysis

Extraction of soluble carbohydrates

The frozen tuber samples were individually ground ina blender with 983090983088 mL of 983095983088 HPLC-grade ethanol and

Condiciones de almacenamiento

En la cosecha los tubeacuterculos de cuatro plantas por unidadexperimental seleccionadas al azar fueron colocados enbolsas de papel para ser almacenados en laboratorio conluz ambiental bajo las condiciones de temperatura yhumedad relativa que se muestran en el Cuadro 983089 hastaque mostraron el maacuteximo desarrollo de brote

Se consideroacute que los tubeacuterculos contenidos en lasbolsas de papel dejaron la condicioacuten de reposo cuandoen 983093983088 de estos el brote principal midioacute 983092 plusmn 983090 mm Lalongitud (mm) y peso seco (mg) del brote se registroacuteen cinco tubeacuterculos elegidos al azar de la unidadexperimental en dominancia apical (un solo brote)brotacioacuten muacuteltiple (maacutes de un brote) y senescencia (maacutesde un brote y peacuterdida de turgencia de los tubeacuterculos)En cada ocasioacuten se midioacute el brote de mayor longitud ya partir de la brotacioacuten muacuteltiple se consideraron todoslos brotes para medir el peso seco

Cuantificacioacuten de carbohidratos solubles por croma-tografiacutea liacutequida de alta resolucioacuten (HPLC)

Obtencioacuten de la muestra

Durante el periodo de almacenamiento se realizaroncuatro muestreos el primero en condiciones dereposo y los otros tres correspondieron a las edadesfisioloacutegicas de dominancia apical brotacioacuten muacuteltiple

Table 983090 Seed-tuber weight at different physiological states of four potato varieties

Cuadro 983090 Peso de semilla-tubeacuterculo en diferentes estados fisioloacutegicos de cuatro variedades de papa

Seed-tuber hysiological state frasl Estado fisioloacutegico de semilla-tubeacuterculo

Variety frasl

VariedadDormancy frasl Reposo

Apical dominance frasl

Dominancia apical

Multiple sprouting frasl

Brotacioacuten muacuteltiple

Senescence frasl

Senescencia

DAHz frasl

DDCz

Weight y (g) frasl

Peso y (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

Alpha 983090983090 983089983097983089 983089983089983090 983089983096983090 983089983092983096 983089983094983091 983090983090983091 983089983093983088

Atlantic 983090983090 983089983092983090 983089983092983095 983089983093983089 983090983089983094 983089983092983093 983090983096983088 983089983091983088

Gigant 983090983090 983090983088983093 983095983095 983090983091983096 983097983096 983090983090983096 983090983090983091 983090983088983088

Vivaldi 983090983090 983089983096983096 983097983096 983089983092983094 983089983092983096 983089983092983097 983090983090983091 983089983091983088

Mean fraslMedia

983090983090 983089983096983089 983089983088983097 983089983095983097 983089983093983091 983089983095983089 983090983091983095 983089983093983090

zDDC days after harvest y average value of two tubers

zDDC diacuteas despueacutes de la cosecha y valor promedio de dos tubeacuterculos

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brought to boiling temperature for 983093 min The extract was decanted and a second extraction was made fromthe residue with 983089983088 mL of ethanol at the end theextracts were mixed Subsequently the extract wascentrifuged for 983089983093 min at 983092983088983088983088 x g in a HERLE modelZ983090983091983088A centrifuge (Labnet International Inc Edison NJ

USA) filtered with Whatman No 983092 paper and mannitol was added at a concentration of 983092 mgmiddotmL-983089 as internal

standard Finally the extract was gauged to 983090983093 mL withthe same kind of solvent used for the extraction

Then 983089983088 mL of the extract were taken and concentratedto dryness with reduced pressure and low temperature(983092983088 degC) in a BUCHI R-983090983089983093 rotary evaporator (Switzerland)The residue was resuspended in 983090 mL of HPLC-grade

water in order to pass it through a column containingion-exchange resins one basic (983088983093 mL of Dowex - 983089 x983096 - Fluka) and the other acidic (983088983093 mL of Dowex - 983093983088W

X983096 - Fluka) at a ratio of 983089983089 (ww) The purified sample was passed through a 983088983092983093-μm nylon acrodisc in orderto collect the sample in a 983089983093 mL vial

HPLC sugar analysis

A Perkin Elmer series 983090983088983088 high-performance liquidchromatography system (Boston Ma USA) consistingof an autosampler quaternary pump degasserrefractive index detector and a column oven wasused The system is operated with TotalChrom version983094983090983089 software A Rezex RCM-monosaccharide Ca+983090 (983096 )column 983091983088 cm in length and 983095983096 mm in diameter wasused As the mobile phase HPLC-grade water was usedThe flow rate was 983088983094 mLmiddotmin-983089 the sample injection

volume 983090983088 μL and the running time 983090983093 min Columntemperature was maintained at 983096983093 degC Pattern curves

were prepared using commercial sucrose glucose andfructose standards (Sigma MN) at concentrations of983088983093 983089983088 983090983093 and 983093 mgmiddotmL-983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) At each point of the curves it was injected at leastthree times to obtain the data of the area associated

with each concentration area using this informationthe regression equations for each sugar were obtained

Analyses of variance and regression were performed with the data obtained Also Tukeyrsquos test was appliedusing the Statistical Analysis System package (SAS 983090983088983088983092)

Results and discussion

Temperature

The air temperature fluctuated during the growingseason (Figure 983089) with the maximum varying between983089983095 and 983091983094 degC Temperatures above 983090983091 degC were recordedfrom 983096 to 983093983092 days after emergence and coincided

with the vegetative (VS) and initial reproductive (RS983089)stages and half of the final reproductive stage (RS 983090)

y senescencia del tubeacuterculo de cada variedad En cadaocasioacuten se tomoacute un tubeacuterculo contenido en las bolsasde papel el de mayor tamantildeo en dos repeticiones conpesos promedio entre 983089983093983090 y 983089983096983089 g (Cuadro 983090) del que seobtuvo la muestra

De la parte central de cada tubeacuterculo entre el extremobasal y apical se extrajo una muestra de 983093 plusmn 983089 g que se

colocoacute en nitroacutegeno liacutequido (-983089983097983094 ordmC) hasta su anaacutelisis

Extraccioacuten de carbohidratos solubles

Las muestras congeladas de tubeacuterculo fueron molidasen licuadora individualmente con 983090983088 mL de etanolgrado HPLC a 983095983088 y llevadas a temperatura deebullicioacuten durante 983093 min El extracto se decantoacute yal residuo se le realizoacute una segunda extraccioacuten con983089983088 mL de etanol al final los extractos se mezclaronPosteriormente el extracto se centrifugoacute por 983089983093 mina 983092983088983088983088 x g en una centriacutefuga HERLE modelo Z983090983091983088A(Labnet International Inc Edison NJ USA) se filtroacute

con papel Whatman nuacutemero 983092 y se agregoacute manitol enconcentracioacuten de 983092 mgmiddotmL-983089 como estaacutendar internoFinalmente el extracto se aforoacute a 983090983093 mL con el mismotipo de solvente empleado para la extraccioacuten

Del extracto aforado se tomaron 983089983088 mL paraconcentrar a sequedad con presioacuten reducida y bajatemperatura (983092983088 ordmC) en un rotoevaporador BUCHIR-983090983089983093 (Switzerland) El residuo se resuspendioacute en 983090 mLde agua grado HPLC para pasarlo por una columnaque conteniacutea resinas intercambiadoras de iones unabaacutesica (983088983093 mL de Dowex - 983089 x 983096 - Fluka) y la otra aacutecida(983088983093 mL de Dowex - 983093983088W X983096 - Fluka) en proporcioacuten 983089983089(pp) La muestra purificada se pasoacute por un acrodiscode nylon de 983088983092983093 μm con el fin de colectar la muestraen un vial de 983089983093 mL

Anaacutelisis de azuacutecares en HPLC

Se utilizoacute un equipo de cromatografiacutea liacutequida de altaresolucioacuten serie 983090983088983088 Perkin Elmer (Boston Ma USA)compuesto por un auto muestreador bomba cuaternariadegasificador detector de iacutendice de refraccioacuten y hornopara columnas El sistema se operoacute con el programacomputacional TotalChrom versioacuten 983094983090983089 Se usoacute una

columna Rezex RCM-monosaccharide Ca+983090

(983096 ) de 983091983088cm de longitud y 983095983096 mm de diaacutemetro Como fase moacutevilse empleoacute agua grado HPLC La velocidad de flujo fuede 983088983094 mLmiddotmin-983089 el volumen de muestra inyectado de983090983088 μL y el tiempo de corrida 983090983093 min La temperatura dela columna se mantuvo a 983096983093 ordmC Se elaboraron curvaspatroacuten usando estaacutendares comerciales de sacarosaglucosa y fructosa (Sigma MN) en concentraciones de983088983093 983089983088 983090983093 y 983093 mgmiddotmL -983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) En cada punto de las curvas se inyectoacute al menostres veces para obtener los datos del aacuterea asociada a

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The minimum ranged between 983095 and 983089983094 degC and eventhough in some periods it was less than 983089983088 degC it did notaffect plant growth and yield since it is considered thatpotato tolerates low temperatures with 983094 degC being itsbase temperature (Cao amp Tibbitts 983089983097983097983093)

The mean temperature was 983089983097 degC similar to thatreported by Cao and Tibbitts (983089983097983097983092) to obtain high

biomass and tuber production (983090983088 degC) Struik Haverkort Vreugdengil Bus and Dankert (983089983097983097983088) stress theimportance of high temperatures during RS 983089 as theypromote stolon branching an important condition for

yield because the number of tubers formed is closelyrelated to the number of stolons (Haverkort Van De

Waart amp Bodlaender 983089983097983097983088)

Crop Phenology

In general there were differential genotypic effects ( P le983088983088983093) in crop phenology The variety Alpha required moreGDD to reach the stolon (983089983094983095) tuberization (983090983094983089) and tuber

physiological maturity (983089983093983092) stages of the tuber its lifecycle was significantly longer than in the other varieties

Vivaldi was the variety that reached maturity the fastestbecause it needed 983089983093 fewer GDD than Alpha (Table 983091)

cada concentracioacuten con esta informacioacuten se obtuvieronlas ecuaciones de regresioacuten para cada azuacutecar

Con los datos obtenidos se realizaron un anaacutelisis de varianza y de regresioacuten ademaacutes se aplicaron pruebas decomparacioacuten de medias de Tukey para ello se empleoacuteel paquete Statistical Analysis System (SAS 983090983088983088983092)

Resultados y discusioacuten

Temperatura

La temperatura del aire fue fluctuante durante el ciclode cultivo (Figura 983089) la maacutexima varioacute entre 983089983095 y 983091983094 ordmCSuperiores a 983090983091 ordmC se registraron desde los 983096 hasta los983093983092 diacuteas despueacutes de la emergencia y coincidieron con laetapa vegetativa (EV) reproductiva inicial (ER 983089) y mitadde la reproductiva final (ER 983090) La miacutenima osciloacute entre 983095

y 983089983094 ordmC y a pesar de que en algunos periodos fue inferiora 983089983088 ordmC no afectoacute al crecimiento y rendimiento de laplanta ya que se considera que la papa resiste bajas

temperaturas siendo su base 983094 ordmC (Cao amp Tibbitts 983089983097983097983093)

La temperatura media fue de 983089983097 ordmC similar a lareportada por Cao y Tibbitts (983089983097983097983092) para obtener alta

Figure 983089 Air temperature recorded during the potato growing season under field conditions San Lorenzo

Cuapiaxtla Tlax Spring-Summer Cycle 983090983088983088983096 The arrows indicate the phases of emergence (E) stolon

initiation (SI) tuberization (T) and tuber physiological maturity (TM) and the intervals between them

at the vegetative (VS) initial reproductive (RS 983089) and final reproductive (RS 983090) stages Average values of

four varieties

Figura 983089 Temperatura del aire registrada durante el ciclo de cultivo de papa en condiciones de campo San

Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096 Las flechas indican las fases de emergencia

(E) inicio de estoloneo (ES) tuberizacioacuten (T) y madurez fisioloacutegica del tubeacuterculo (MT) y los intervalos

entre estas a las etapas vegetativa (EV) reproductiva inicial (ER 983089) y reproductiva final (ER 983090) Valores

promedio de cuatro variedades

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The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

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Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

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Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

change patterns affect growth and tuberization inpotatoes Journal of the American Society for Horticultural

Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 318

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

The results of studies on the role of sugars intuber sprouting are ambiguous Dogras Siomosand Psomakelis (983089983097983096983097) observed an increase in theconcentration of total and reducing sugars in sproutedtubers of the variety Spunta whereas in cv Sebagothe content remained constant Debast et al (983090983088983089983089)indicate that the transport of sucrose to the tuberbuds is a prerequisite to induce sprouting and during

this stage the soluble sugar content decreases untilthe tuber sprouts reach approximately 983089 g dry matter(Viola et al 983090983088983088983095)

Therefore the objective of this research was to assessthe growth and yield of four potato varieties and todetermine the sugar content in tubers at differentphysiological ages and the effect these sugars have onseed-tuber sprouting

Materials and methods

This research was conducted in the municipality of San

Lorenzo Cuapiaxtla Tlaxcala (983089983097ordm 983089983096rsquo N 983097983095ordm 983092983094rsquo W and983090983092983089983088 masl) under irrigated conditions The regionrsquosclimate classification is C (Wo) (w) b (irsquo) C (Wo) (Garciacutea983089983097983096983096) and it has regosol-type soil

Four varieties were evaluated Alpha Atlantic Gigantand Vivaldi At planting minitubers of 983089983094 to 983090983096 mmin diameter in a state of apical dominance were usedThe experimental unit consisted of a row of 983093 m longand 983088983097983088 m wide with 983088983090983093 m spacing between plantsin each row 983090983088 minitubers were deposited at a depthof 983089983088 cm

At planting and 983090983088 days after emergence theminitubers were fertilized with the formula 983090983093983088-983094983088-983091983093983088-983097983089-983094983091 (N P K Ca and Mg) During the growingseason it was spray irrigated according to crop waterrequirements From emergence to tuber physiologicalmaturity the maximum and minimum air temperature(degC) was recorded with a Taylorreg model 983093983092983093983096 mercurythermometer using these data the mean temperature

was obtained

A randomized complete block experimental design with four replications was used and the experimental

unit consisted of four rows

From emergence the number of growing degree-days(GDD) accumulated at the beginning of each phaseand during each phenological stage of the crop wasdetermined with the classical residual method and983094 degC base temperature (Cao amp Tibbitts 983089983097983097983093) Theemergence phase (E) occurred when 983093983088 of the plantsof each experimental unit had emerged The stoloninitiation (SI) phase began when the stolon appearedon the main stem Tuber initiation or growth (T) was

de 983089983090983096 de azuacutecares totales y 983088983093983093 de azuacutecaresreductores En otros estudios Rodriacuteguez-Saona y

Wrolstad (983089983097983097983095) reportaron valores de 983088983089983091 desacarosa 983088983088983092 de glucosa y 983088983088983091 de fructosamientras que Sabba et al (983090983088983088983095) indican valores de983088983089983089 de sacarosa y 983088983088983093 de glucosa en tubeacuterculos enestado de reposo

Los resultados de estudios sobre el papel de losazuacutecares en la brotacioacuten de los tubeacuterculos sonpoco claros Dogras Siomos y Psomakelis (983089983097983096983097)observaron incremento en la concentracioacuten deazuacutecares totales y reductores en tubeacuterculos brotadosde la variedad Spunta en tanto que en cv Sebagoel contenido se mantuvo constante Debast et al(983090983088983089983089) indican que el transporte de sacarosa a las

yemas del tubeacuterculo es prerrequisito para inducir labrotacioacuten y durante eacutesta el contenido de azuacutecaressolubles disminuye hasta que los brotes del tubeacuterculoalcanzan aproximadamente 983089 g de materia seca(Viola et al 983090983088983088983095)

Por lo anterior la presente investigacioacuten tuvo comoobjetivo evaluar el crecimiento y rendimiento de cuatro

variedades de papa determinar el contenido de azuacutecaresen tubeacuterculos con diferente edad fisioloacutegica y el efectoque tienen estos en la brotacioacuten de la semilla-tubeacuterculo

Materiales y meacutetodos

La presente investigacioacuten se realizoacute en el municipiode San Lorenzo Cuapiaxtla Tlaxcala (983089983097ordm 983089983096rsquo N 983097983095ordm 983092983094rsquoO y 983090983092983089983088 msnm) bajo condiciones de riego La regioacutenposee clima C (Wo) (w) b (irsquo) C (Wo) (Garciacutea 983089983097983096983096) ysuelo de tipo regosol

Se evaluaron las variedades Alpha Atlantic Gigant y Vivaldi En la siembra se usaron minitubeacuterculos de 983089983094a 983090983096 mm de diaacutemetro en estado de dominancia apicalla unidad experimental consistioacute en un surco de 983093 m delongitud con 983088983097983088 m de ancho y 983088983090983093 m entre plantasen cada surco se depositaron 983090983088 minitubeacuterculos a 983089983088cm de profundidad

En la siembra y 983090983088 diacuteas despueacutes de la emergencia sefertilizoacute con la foacutermula 983090983093983088-983094983088-983091983093983088-983097983089-983094983091 (N P K Ca y

Mg) Durante el ciclo de cultivo se irrigoacute por aspersioacutende acuerdo con las necesidades hiacutedricas del mismoDesde la emergencia y hasta la madurez fisioloacutegicadel tubeacuterculo se registroacute la temperatura maacutexima ymiacutenima del aire (ordmC) con un termoacutemetro de columna demercurio marca Taylorreg modelo 983093983092983093983096 con esos datos seobtuvo la temperatura media

El disentildeo experimental fue bloques completos al azarcon cuatro repeticiones y la unidad experimentalestuvo constituida por cuatro surcos

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identified as the time at which the apex of the stolonthickened Tuber physiological maturity (TM) was

visually identified by the loss of the foliagersquos greencolor or onset of senescence

The vegetative stage (VS) was considered as the timefrom emergence to the beginning of the stolonphase during which shoot growth and root system

establishment occur The initial reproductive stage (RS983089) includes the interval between the onset of the stolonstage and tuberization and is characterized by stolongeneration and growth The final reproductive stage(RS 983090) runs from the beginning of tuberization to tuberphysiological maturity where tuber growth occurred(Morales-Fernaacutendez et al 983090983088983089983089)

At harvest the number (NCTP) and weight (WCTP g) ofcommercial tubers per plant (with a greater than 983090983088-mmdiameter) were quantified as were the number (NNCTP)and weight (WNCTP g) of non-commercial tubers perplant (with a 983090983088-mm diameter) characterized by being

deformed cracked and green The total number of tubersper plant (TNTP) was obtained by adding NCTP and NNCTPThe total weight of tubers per plant (TWTP g) was the sumof WCTP plus WNCTP The average tuber weight per plant(ATWP) was obtained by dividing TWTP by TNTP

Storage conditions

At harvest the tubers of four randomly-selected plantsper experimental unit were placed in paper bags to belater stored in the laboratory with ambient light underthe temperature and relative humidity conditions shownin Table 983089 until showing maximum sprout developmentIt was considered that the tubers contained in thepaper bags left dormancy when in 983093983088 of them the mainsprout measured 983092 plusmn 983090 mm Sprout length (mm) anddry weight (mg) were recorded in five tubers selected

A partir de la emergencia se determinoacute el nuacutemerode grados diacutea (GD) acumulados al inicio de cada fase

y durante cada etapa fenoloacutegica del cultivo con elmeacutetodo residual claacutesico y la temperatura base de 983094 ordmC (Caoamp Tibbitts 983089983097983097983093) La fase de emergencia (E) ocurrioacute cuando983093983088 de las plantas de cada unidad experimental habiacuteanemergido El inicio de estoloneo (ES) fue cuando aparecioacuteel estoloacuten en el tallo principal El inicio de tuberizacioacuten

o de crecimiento del tubeacuterculo (T) se identificoacute como elmomento en que engrosoacute el aacutepice del estoloacuten La madurezfisioloacutegica del tubeacuterculo (MT) se identificoacute visualmentepor la peacuterdida del color verde del follaje o inicio de lasenescencia de eacuteste

La etapa vegetativa (EV) se consideroacute como el periodotranscurrido desde la emergencia hasta el inicio deestoloneo durante el cual ocurre el crecimiento de laparte aeacuterea y el establecimiento del sistema radicalLa etapa reproductiva inicial (ER 983089) incluyoacute el intervaloentre el inicio de estoloneo y el de tuberizacioacuten y secaracterizoacute por la generacioacuten y crecimiento de estolones

La etapa reproductiva final (ER 983090) va desde el inicio de latuberizacioacuten hasta la madurez fisioloacutegica del tubeacuterculodonde ocurrioacute el crecimiento de este oacutergano (Morales-Fernaacutendez et al 983090983088983089983089)

Durante la cosecha se cuantificoacute el nuacutemero detubeacuterculos por planta (NTCP) y peso de tubeacuterculos porplanta (PTCP g) de tubeacuterculos comerciales por planta(con diaacutemetro mayor de 983090983088 mm) nuacutemero (NTNCP)

y peso (PTNCP g) de tubeacuterculos no comerciales porplanta (con diaacutemetro menor de 983090983088 mm) deformesrajados y verdes El nuacutemero total de tubeacuterculos porplanta (NTTP) que se obtuvo al sumar NTCP maacutesNTNCP El peso total de tubeacuterculos por planta (PTTPg) fue la suma de PTCP maacutes PTNCP El peso promediode tubeacuterculo por planta (PPTP) se obtuvo al dividirPTTP entre NTTP

Table 983089 Air temperature and relative humidity recorded during storage of potato seed-tubers after harvest

Cuadro 983089 Temperatura del aire y humedad relativa registradas durante el almacenamiento de la semilla-

tubeacuterculo de papa despueacutes de la cosecha

ELEMENT frasl ELEMENTO

983090983088983088983096 983090983088983088983097

AGO frasl

AUGSEP OCT NOV

DIC frasl

DECENE FEB MAR

ABR frasl

APR MAY JUN

Temperature (ordmC) frasl Temperatura (ordmC)

Maximum frasl Maacutexima 983090983094 983090983092 983090983089 983089983095 983089983094 983089983093 983089983095 983090983089 983090983092 983090983094 983090983095

Minimum frasl Miacutenima 983089983096 983089983096 983089983094 983089983090 983089983090 983089983089 983089983092 983089983092 983089983095 983090983089 983090983088

Mean frasl Media 983090983090 983090983089 983089983096 983089983092 983089983092 983089983091 983089983093 983089983095 983090983088 983090983091 983090983091

Humedad relativa () frasl

Relative humidity ()

Maximum frasl Maacutexima 983092983090 983092983092 983091983093 983091983088 983091983088 983091983089 983091983091 983091983090 983091983092 983091983094 983092983094

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at random from the experimental unit during apicaldominance (a single sprout) multiple sprouting (morethan one sprout) and senescence (more than onesprout and turgor loss in tubers) On each occasion the longest sprout was measured and from multiplesprouting all sprouts were measured for dry weight

Quantification of soluble carbohydrates by high-

performance liquid chromatography (HPLC)

Sample collection

Four samplings were performed during the storageperiod the first during dormancy and the other threeduring the physiological ages of apical dominancemultiple sprouting and senescence of the tuber ofeach variety On each occasion a tuber contained in thepaper bags was taken and a sample obtained from it

with two replications being made for the larger tubers Average tuber weights for the sampling process rangedbetween 983089983093983090 and 983089983096983089 g (Table 983090)

From the central part of each tuber between the basaland apical end a sample of 983093 plusmn 983089 g was extracted andplaced in liquid nitrogen (-983089983097983094 degC) until analysis

Extraction of soluble carbohydrates

The frozen tuber samples were individually ground ina blender with 983090983088 mL of 983095983088 HPLC-grade ethanol and

Condiciones de almacenamiento

En la cosecha los tubeacuterculos de cuatro plantas por unidadexperimental seleccionadas al azar fueron colocados enbolsas de papel para ser almacenados en laboratorio conluz ambiental bajo las condiciones de temperatura yhumedad relativa que se muestran en el Cuadro 983089 hastaque mostraron el maacuteximo desarrollo de brote

Se consideroacute que los tubeacuterculos contenidos en lasbolsas de papel dejaron la condicioacuten de reposo cuandoen 983093983088 de estos el brote principal midioacute 983092 plusmn 983090 mm Lalongitud (mm) y peso seco (mg) del brote se registroacuteen cinco tubeacuterculos elegidos al azar de la unidadexperimental en dominancia apical (un solo brote)brotacioacuten muacuteltiple (maacutes de un brote) y senescencia (maacutesde un brote y peacuterdida de turgencia de los tubeacuterculos)En cada ocasioacuten se midioacute el brote de mayor longitud ya partir de la brotacioacuten muacuteltiple se consideraron todoslos brotes para medir el peso seco

Cuantificacioacuten de carbohidratos solubles por croma-tografiacutea liacutequida de alta resolucioacuten (HPLC)

Obtencioacuten de la muestra

Durante el periodo de almacenamiento se realizaroncuatro muestreos el primero en condiciones dereposo y los otros tres correspondieron a las edadesfisioloacutegicas de dominancia apical brotacioacuten muacuteltiple

Table 983090 Seed-tuber weight at different physiological states of four potato varieties

Cuadro 983090 Peso de semilla-tubeacuterculo en diferentes estados fisioloacutegicos de cuatro variedades de papa

Seed-tuber hysiological state frasl Estado fisioloacutegico de semilla-tubeacuterculo

Variety frasl

VariedadDormancy frasl Reposo

Apical dominance frasl

Dominancia apical

Multiple sprouting frasl

Brotacioacuten muacuteltiple

Senescence frasl

Senescencia

DAHz frasl

DDCz

Weight y (g) frasl

Peso y (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

Alpha 983090983090 983089983097983089 983089983089983090 983089983096983090 983089983092983096 983089983094983091 983090983090983091 983089983093983088

Atlantic 983090983090 983089983092983090 983089983092983095 983089983093983089 983090983089983094 983089983092983093 983090983096983088 983089983091983088

Gigant 983090983090 983090983088983093 983095983095 983090983091983096 983097983096 983090983090983096 983090983090983091 983090983088983088

Vivaldi 983090983090 983089983096983096 983097983096 983089983092983094 983089983092983096 983089983092983097 983090983090983091 983089983091983088

Mean fraslMedia

983090983090 983089983096983089 983089983088983097 983089983095983097 983089983093983091 983089983095983089 983090983091983095 983089983093983090

zDDC days after harvest y average value of two tubers

zDDC diacuteas despueacutes de la cosecha y valor promedio de dos tubeacuterculos

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brought to boiling temperature for 983093 min The extract was decanted and a second extraction was made fromthe residue with 983089983088 mL of ethanol at the end theextracts were mixed Subsequently the extract wascentrifuged for 983089983093 min at 983092983088983088983088 x g in a HERLE modelZ983090983091983088A centrifuge (Labnet International Inc Edison NJ

USA) filtered with Whatman No 983092 paper and mannitol was added at a concentration of 983092 mgmiddotmL-983089 as internal

standard Finally the extract was gauged to 983090983093 mL withthe same kind of solvent used for the extraction

Then 983089983088 mL of the extract were taken and concentratedto dryness with reduced pressure and low temperature(983092983088 degC) in a BUCHI R-983090983089983093 rotary evaporator (Switzerland)The residue was resuspended in 983090 mL of HPLC-grade

water in order to pass it through a column containingion-exchange resins one basic (983088983093 mL of Dowex - 983089 x983096 - Fluka) and the other acidic (983088983093 mL of Dowex - 983093983088W

X983096 - Fluka) at a ratio of 983089983089 (ww) The purified sample was passed through a 983088983092983093-μm nylon acrodisc in orderto collect the sample in a 983089983093 mL vial

HPLC sugar analysis

A Perkin Elmer series 983090983088983088 high-performance liquidchromatography system (Boston Ma USA) consistingof an autosampler quaternary pump degasserrefractive index detector and a column oven wasused The system is operated with TotalChrom version983094983090983089 software A Rezex RCM-monosaccharide Ca+983090 (983096 )column 983091983088 cm in length and 983095983096 mm in diameter wasused As the mobile phase HPLC-grade water was usedThe flow rate was 983088983094 mLmiddotmin-983089 the sample injection

volume 983090983088 μL and the running time 983090983093 min Columntemperature was maintained at 983096983093 degC Pattern curves

were prepared using commercial sucrose glucose andfructose standards (Sigma MN) at concentrations of983088983093 983089983088 983090983093 and 983093 mgmiddotmL-983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) At each point of the curves it was injected at leastthree times to obtain the data of the area associated

with each concentration area using this informationthe regression equations for each sugar were obtained

Analyses of variance and regression were performed with the data obtained Also Tukeyrsquos test was appliedusing the Statistical Analysis System package (SAS 983090983088983088983092)

Results and discussion

Temperature

The air temperature fluctuated during the growingseason (Figure 983089) with the maximum varying between983089983095 and 983091983094 degC Temperatures above 983090983091 degC were recordedfrom 983096 to 983093983092 days after emergence and coincided

with the vegetative (VS) and initial reproductive (RS983089)stages and half of the final reproductive stage (RS 983090)

y senescencia del tubeacuterculo de cada variedad En cadaocasioacuten se tomoacute un tubeacuterculo contenido en las bolsasde papel el de mayor tamantildeo en dos repeticiones conpesos promedio entre 983089983093983090 y 983089983096983089 g (Cuadro 983090) del que seobtuvo la muestra

De la parte central de cada tubeacuterculo entre el extremobasal y apical se extrajo una muestra de 983093 plusmn 983089 g que se

colocoacute en nitroacutegeno liacutequido (-983089983097983094 ordmC) hasta su anaacutelisis

Extraccioacuten de carbohidratos solubles

Las muestras congeladas de tubeacuterculo fueron molidasen licuadora individualmente con 983090983088 mL de etanolgrado HPLC a 983095983088 y llevadas a temperatura deebullicioacuten durante 983093 min El extracto se decantoacute yal residuo se le realizoacute una segunda extraccioacuten con983089983088 mL de etanol al final los extractos se mezclaronPosteriormente el extracto se centrifugoacute por 983089983093 mina 983092983088983088983088 x g en una centriacutefuga HERLE modelo Z983090983091983088A(Labnet International Inc Edison NJ USA) se filtroacute

con papel Whatman nuacutemero 983092 y se agregoacute manitol enconcentracioacuten de 983092 mgmiddotmL-983089 como estaacutendar internoFinalmente el extracto se aforoacute a 983090983093 mL con el mismotipo de solvente empleado para la extraccioacuten

Del extracto aforado se tomaron 983089983088 mL paraconcentrar a sequedad con presioacuten reducida y bajatemperatura (983092983088 ordmC) en un rotoevaporador BUCHIR-983090983089983093 (Switzerland) El residuo se resuspendioacute en 983090 mLde agua grado HPLC para pasarlo por una columnaque conteniacutea resinas intercambiadoras de iones unabaacutesica (983088983093 mL de Dowex - 983089 x 983096 - Fluka) y la otra aacutecida(983088983093 mL de Dowex - 983093983088W X983096 - Fluka) en proporcioacuten 983089983089(pp) La muestra purificada se pasoacute por un acrodiscode nylon de 983088983092983093 μm con el fin de colectar la muestraen un vial de 983089983093 mL

Anaacutelisis de azuacutecares en HPLC

Se utilizoacute un equipo de cromatografiacutea liacutequida de altaresolucioacuten serie 983090983088983088 Perkin Elmer (Boston Ma USA)compuesto por un auto muestreador bomba cuaternariadegasificador detector de iacutendice de refraccioacuten y hornopara columnas El sistema se operoacute con el programacomputacional TotalChrom versioacuten 983094983090983089 Se usoacute una

columna Rezex RCM-monosaccharide Ca+983090

(983096 ) de 983091983088cm de longitud y 983095983096 mm de diaacutemetro Como fase moacutevilse empleoacute agua grado HPLC La velocidad de flujo fuede 983088983094 mLmiddotmin-983089 el volumen de muestra inyectado de983090983088 μL y el tiempo de corrida 983090983093 min La temperatura dela columna se mantuvo a 983096983093 ordmC Se elaboraron curvaspatroacuten usando estaacutendares comerciales de sacarosaglucosa y fructosa (Sigma MN) en concentraciones de983088983093 983089983088 983090983093 y 983093 mgmiddotmL -983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) En cada punto de las curvas se inyectoacute al menostres veces para obtener los datos del aacuterea asociada a

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The minimum ranged between 983095 and 983089983094 degC and eventhough in some periods it was less than 983089983088 degC it did notaffect plant growth and yield since it is considered thatpotato tolerates low temperatures with 983094 degC being itsbase temperature (Cao amp Tibbitts 983089983097983097983093)

The mean temperature was 983089983097 degC similar to thatreported by Cao and Tibbitts (983089983097983097983092) to obtain high

biomass and tuber production (983090983088 degC) Struik Haverkort Vreugdengil Bus and Dankert (983089983097983097983088) stress theimportance of high temperatures during RS 983089 as theypromote stolon branching an important condition for

yield because the number of tubers formed is closelyrelated to the number of stolons (Haverkort Van De

Waart amp Bodlaender 983089983097983097983088)

Crop Phenology

In general there were differential genotypic effects ( P le983088983088983093) in crop phenology The variety Alpha required moreGDD to reach the stolon (983089983094983095) tuberization (983090983094983089) and tuber

physiological maturity (983089983093983092) stages of the tuber its lifecycle was significantly longer than in the other varieties

Vivaldi was the variety that reached maturity the fastestbecause it needed 983089983093 fewer GDD than Alpha (Table 983091)

cada concentracioacuten con esta informacioacuten se obtuvieronlas ecuaciones de regresioacuten para cada azuacutecar

Con los datos obtenidos se realizaron un anaacutelisis de varianza y de regresioacuten ademaacutes se aplicaron pruebas decomparacioacuten de medias de Tukey para ello se empleoacuteel paquete Statistical Analysis System (SAS 983090983088983088983092)

Resultados y discusioacuten

Temperatura

La temperatura del aire fue fluctuante durante el ciclode cultivo (Figura 983089) la maacutexima varioacute entre 983089983095 y 983091983094 ordmCSuperiores a 983090983091 ordmC se registraron desde los 983096 hasta los983093983092 diacuteas despueacutes de la emergencia y coincidieron con laetapa vegetativa (EV) reproductiva inicial (ER 983089) y mitadde la reproductiva final (ER 983090) La miacutenima osciloacute entre 983095

y 983089983094 ordmC y a pesar de que en algunos periodos fue inferiora 983089983088 ordmC no afectoacute al crecimiento y rendimiento de laplanta ya que se considera que la papa resiste bajas

temperaturas siendo su base 983094 ordmC (Cao amp Tibbitts 983089983097983097983093)

La temperatura media fue de 983089983097 ordmC similar a lareportada por Cao y Tibbitts (983089983097983097983092) para obtener alta

Figure 983089 Air temperature recorded during the potato growing season under field conditions San Lorenzo

Cuapiaxtla Tlax Spring-Summer Cycle 983090983088983088983096 The arrows indicate the phases of emergence (E) stolon

initiation (SI) tuberization (T) and tuber physiological maturity (TM) and the intervals between them

at the vegetative (VS) initial reproductive (RS 983089) and final reproductive (RS 983090) stages Average values of

four varieties

Figura 983089 Temperatura del aire registrada durante el ciclo de cultivo de papa en condiciones de campo San

Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096 Las flechas indican las fases de emergencia

(E) inicio de estoloneo (ES) tuberizacioacuten (T) y madurez fisioloacutegica del tubeacuterculo (MT) y los intervalos

entre estas a las etapas vegetativa (EV) reproductiva inicial (ER 983089) y reproductiva final (ER 983090) Valores

promedio de cuatro variedades

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The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

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Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

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Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

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Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

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Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

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acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

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M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

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development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

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7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 418

983089983091983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

identified as the time at which the apex of the stolonthickened Tuber physiological maturity (TM) was

visually identified by the loss of the foliagersquos greencolor or onset of senescence

The vegetative stage (VS) was considered as the timefrom emergence to the beginning of the stolonphase during which shoot growth and root system

establishment occur The initial reproductive stage (RS983089) includes the interval between the onset of the stolonstage and tuberization and is characterized by stolongeneration and growth The final reproductive stage(RS 983090) runs from the beginning of tuberization to tuberphysiological maturity where tuber growth occurred(Morales-Fernaacutendez et al 983090983088983089983089)

At harvest the number (NCTP) and weight (WCTP g) ofcommercial tubers per plant (with a greater than 983090983088-mmdiameter) were quantified as were the number (NNCTP)and weight (WNCTP g) of non-commercial tubers perplant (with a 983090983088-mm diameter) characterized by being

deformed cracked and green The total number of tubersper plant (TNTP) was obtained by adding NCTP and NNCTPThe total weight of tubers per plant (TWTP g) was the sumof WCTP plus WNCTP The average tuber weight per plant(ATWP) was obtained by dividing TWTP by TNTP

Storage conditions

At harvest the tubers of four randomly-selected plantsper experimental unit were placed in paper bags to belater stored in the laboratory with ambient light underthe temperature and relative humidity conditions shownin Table 983089 until showing maximum sprout developmentIt was considered that the tubers contained in thepaper bags left dormancy when in 983093983088 of them the mainsprout measured 983092 plusmn 983090 mm Sprout length (mm) anddry weight (mg) were recorded in five tubers selected

A partir de la emergencia se determinoacute el nuacutemerode grados diacutea (GD) acumulados al inicio de cada fase

y durante cada etapa fenoloacutegica del cultivo con elmeacutetodo residual claacutesico y la temperatura base de 983094 ordmC (Caoamp Tibbitts 983089983097983097983093) La fase de emergencia (E) ocurrioacute cuando983093983088 de las plantas de cada unidad experimental habiacuteanemergido El inicio de estoloneo (ES) fue cuando aparecioacuteel estoloacuten en el tallo principal El inicio de tuberizacioacuten

o de crecimiento del tubeacuterculo (T) se identificoacute como elmomento en que engrosoacute el aacutepice del estoloacuten La madurezfisioloacutegica del tubeacuterculo (MT) se identificoacute visualmentepor la peacuterdida del color verde del follaje o inicio de lasenescencia de eacuteste

La etapa vegetativa (EV) se consideroacute como el periodotranscurrido desde la emergencia hasta el inicio deestoloneo durante el cual ocurre el crecimiento de laparte aeacuterea y el establecimiento del sistema radicalLa etapa reproductiva inicial (ER 983089) incluyoacute el intervaloentre el inicio de estoloneo y el de tuberizacioacuten y secaracterizoacute por la generacioacuten y crecimiento de estolones

La etapa reproductiva final (ER 983090) va desde el inicio de latuberizacioacuten hasta la madurez fisioloacutegica del tubeacuterculodonde ocurrioacute el crecimiento de este oacutergano (Morales-Fernaacutendez et al 983090983088983089983089)

Durante la cosecha se cuantificoacute el nuacutemero detubeacuterculos por planta (NTCP) y peso de tubeacuterculos porplanta (PTCP g) de tubeacuterculos comerciales por planta(con diaacutemetro mayor de 983090983088 mm) nuacutemero (NTNCP)

y peso (PTNCP g) de tubeacuterculos no comerciales porplanta (con diaacutemetro menor de 983090983088 mm) deformesrajados y verdes El nuacutemero total de tubeacuterculos porplanta (NTTP) que se obtuvo al sumar NTCP maacutesNTNCP El peso total de tubeacuterculos por planta (PTTPg) fue la suma de PTCP maacutes PTNCP El peso promediode tubeacuterculo por planta (PPTP) se obtuvo al dividirPTTP entre NTTP

Table 983089 Air temperature and relative humidity recorded during storage of potato seed-tubers after harvest

Cuadro 983089 Temperatura del aire y humedad relativa registradas durante el almacenamiento de la semilla-

tubeacuterculo de papa despueacutes de la cosecha

ELEMENT frasl ELEMENTO

983090983088983088983096 983090983088983088983097

AGO frasl

AUGSEP OCT NOV

DIC frasl

DECENE FEB MAR

ABR frasl

APR MAY JUN

Temperature (ordmC) frasl Temperatura (ordmC)

Maximum frasl Maacutexima 983090983094 983090983092 983090983089 983089983095 983089983094 983089983093 983089983095 983090983089 983090983092 983090983094 983090983095

Minimum frasl Miacutenima 983089983096 983089983096 983089983094 983089983090 983089983090 983089983089 983089983092 983089983092 983089983095 983090983089 983090983088

Mean frasl Media 983090983090 983090983089 983089983096 983089983092 983089983092 983089983091 983089983093 983089983095 983090983088 983090983091 983090983091

Humedad relativa () frasl

Relative humidity ()

Maximum frasl Maacutexima 983092983090 983092983092 983091983093 983091983088 983091983088 983091983089 983091983091 983091983090 983091983092 983091983094 983092983094

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at random from the experimental unit during apicaldominance (a single sprout) multiple sprouting (morethan one sprout) and senescence (more than onesprout and turgor loss in tubers) On each occasion the longest sprout was measured and from multiplesprouting all sprouts were measured for dry weight

Quantification of soluble carbohydrates by high-

performance liquid chromatography (HPLC)

Sample collection

Four samplings were performed during the storageperiod the first during dormancy and the other threeduring the physiological ages of apical dominancemultiple sprouting and senescence of the tuber ofeach variety On each occasion a tuber contained in thepaper bags was taken and a sample obtained from it

with two replications being made for the larger tubers Average tuber weights for the sampling process rangedbetween 983089983093983090 and 983089983096983089 g (Table 983090)

From the central part of each tuber between the basaland apical end a sample of 983093 plusmn 983089 g was extracted andplaced in liquid nitrogen (-983089983097983094 degC) until analysis

Extraction of soluble carbohydrates

The frozen tuber samples were individually ground ina blender with 983090983088 mL of 983095983088 HPLC-grade ethanol and

Condiciones de almacenamiento

En la cosecha los tubeacuterculos de cuatro plantas por unidadexperimental seleccionadas al azar fueron colocados enbolsas de papel para ser almacenados en laboratorio conluz ambiental bajo las condiciones de temperatura yhumedad relativa que se muestran en el Cuadro 983089 hastaque mostraron el maacuteximo desarrollo de brote

Se consideroacute que los tubeacuterculos contenidos en lasbolsas de papel dejaron la condicioacuten de reposo cuandoen 983093983088 de estos el brote principal midioacute 983092 plusmn 983090 mm Lalongitud (mm) y peso seco (mg) del brote se registroacuteen cinco tubeacuterculos elegidos al azar de la unidadexperimental en dominancia apical (un solo brote)brotacioacuten muacuteltiple (maacutes de un brote) y senescencia (maacutesde un brote y peacuterdida de turgencia de los tubeacuterculos)En cada ocasioacuten se midioacute el brote de mayor longitud ya partir de la brotacioacuten muacuteltiple se consideraron todoslos brotes para medir el peso seco

Cuantificacioacuten de carbohidratos solubles por croma-tografiacutea liacutequida de alta resolucioacuten (HPLC)

Obtencioacuten de la muestra

Durante el periodo de almacenamiento se realizaroncuatro muestreos el primero en condiciones dereposo y los otros tres correspondieron a las edadesfisioloacutegicas de dominancia apical brotacioacuten muacuteltiple

Table 983090 Seed-tuber weight at different physiological states of four potato varieties

Cuadro 983090 Peso de semilla-tubeacuterculo en diferentes estados fisioloacutegicos de cuatro variedades de papa

Seed-tuber hysiological state frasl Estado fisioloacutegico de semilla-tubeacuterculo

Variety frasl

VariedadDormancy frasl Reposo

Apical dominance frasl

Dominancia apical

Multiple sprouting frasl

Brotacioacuten muacuteltiple

Senescence frasl

Senescencia

DAHz frasl

DDCz

Weight y (g) frasl

Peso y (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

Alpha 983090983090 983089983097983089 983089983089983090 983089983096983090 983089983092983096 983089983094983091 983090983090983091 983089983093983088

Atlantic 983090983090 983089983092983090 983089983092983095 983089983093983089 983090983089983094 983089983092983093 983090983096983088 983089983091983088

Gigant 983090983090 983090983088983093 983095983095 983090983091983096 983097983096 983090983090983096 983090983090983091 983090983088983088

Vivaldi 983090983090 983089983096983096 983097983096 983089983092983094 983089983092983096 983089983092983097 983090983090983091 983089983091983088

Mean fraslMedia

983090983090 983089983096983089 983089983088983097 983089983095983097 983089983093983091 983089983095983089 983090983091983095 983089983093983090

zDDC days after harvest y average value of two tubers

zDDC diacuteas despueacutes de la cosecha y valor promedio de dos tubeacuterculos

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brought to boiling temperature for 983093 min The extract was decanted and a second extraction was made fromthe residue with 983089983088 mL of ethanol at the end theextracts were mixed Subsequently the extract wascentrifuged for 983089983093 min at 983092983088983088983088 x g in a HERLE modelZ983090983091983088A centrifuge (Labnet International Inc Edison NJ

USA) filtered with Whatman No 983092 paper and mannitol was added at a concentration of 983092 mgmiddotmL-983089 as internal

standard Finally the extract was gauged to 983090983093 mL withthe same kind of solvent used for the extraction

Then 983089983088 mL of the extract were taken and concentratedto dryness with reduced pressure and low temperature(983092983088 degC) in a BUCHI R-983090983089983093 rotary evaporator (Switzerland)The residue was resuspended in 983090 mL of HPLC-grade

water in order to pass it through a column containingion-exchange resins one basic (983088983093 mL of Dowex - 983089 x983096 - Fluka) and the other acidic (983088983093 mL of Dowex - 983093983088W

X983096 - Fluka) at a ratio of 983089983089 (ww) The purified sample was passed through a 983088983092983093-μm nylon acrodisc in orderto collect the sample in a 983089983093 mL vial

HPLC sugar analysis

A Perkin Elmer series 983090983088983088 high-performance liquidchromatography system (Boston Ma USA) consistingof an autosampler quaternary pump degasserrefractive index detector and a column oven wasused The system is operated with TotalChrom version983094983090983089 software A Rezex RCM-monosaccharide Ca+983090 (983096 )column 983091983088 cm in length and 983095983096 mm in diameter wasused As the mobile phase HPLC-grade water was usedThe flow rate was 983088983094 mLmiddotmin-983089 the sample injection

volume 983090983088 μL and the running time 983090983093 min Columntemperature was maintained at 983096983093 degC Pattern curves

were prepared using commercial sucrose glucose andfructose standards (Sigma MN) at concentrations of983088983093 983089983088 983090983093 and 983093 mgmiddotmL-983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) At each point of the curves it was injected at leastthree times to obtain the data of the area associated

with each concentration area using this informationthe regression equations for each sugar were obtained

Analyses of variance and regression were performed with the data obtained Also Tukeyrsquos test was appliedusing the Statistical Analysis System package (SAS 983090983088983088983092)

Results and discussion

Temperature

The air temperature fluctuated during the growingseason (Figure 983089) with the maximum varying between983089983095 and 983091983094 degC Temperatures above 983090983091 degC were recordedfrom 983096 to 983093983092 days after emergence and coincided

with the vegetative (VS) and initial reproductive (RS983089)stages and half of the final reproductive stage (RS 983090)

y senescencia del tubeacuterculo de cada variedad En cadaocasioacuten se tomoacute un tubeacuterculo contenido en las bolsasde papel el de mayor tamantildeo en dos repeticiones conpesos promedio entre 983089983093983090 y 983089983096983089 g (Cuadro 983090) del que seobtuvo la muestra

De la parte central de cada tubeacuterculo entre el extremobasal y apical se extrajo una muestra de 983093 plusmn 983089 g que se

colocoacute en nitroacutegeno liacutequido (-983089983097983094 ordmC) hasta su anaacutelisis

Extraccioacuten de carbohidratos solubles

Las muestras congeladas de tubeacuterculo fueron molidasen licuadora individualmente con 983090983088 mL de etanolgrado HPLC a 983095983088 y llevadas a temperatura deebullicioacuten durante 983093 min El extracto se decantoacute yal residuo se le realizoacute una segunda extraccioacuten con983089983088 mL de etanol al final los extractos se mezclaronPosteriormente el extracto se centrifugoacute por 983089983093 mina 983092983088983088983088 x g en una centriacutefuga HERLE modelo Z983090983091983088A(Labnet International Inc Edison NJ USA) se filtroacute

con papel Whatman nuacutemero 983092 y se agregoacute manitol enconcentracioacuten de 983092 mgmiddotmL-983089 como estaacutendar internoFinalmente el extracto se aforoacute a 983090983093 mL con el mismotipo de solvente empleado para la extraccioacuten

Del extracto aforado se tomaron 983089983088 mL paraconcentrar a sequedad con presioacuten reducida y bajatemperatura (983092983088 ordmC) en un rotoevaporador BUCHIR-983090983089983093 (Switzerland) El residuo se resuspendioacute en 983090 mLde agua grado HPLC para pasarlo por una columnaque conteniacutea resinas intercambiadoras de iones unabaacutesica (983088983093 mL de Dowex - 983089 x 983096 - Fluka) y la otra aacutecida(983088983093 mL de Dowex - 983093983088W X983096 - Fluka) en proporcioacuten 983089983089(pp) La muestra purificada se pasoacute por un acrodiscode nylon de 983088983092983093 μm con el fin de colectar la muestraen un vial de 983089983093 mL

Anaacutelisis de azuacutecares en HPLC

Se utilizoacute un equipo de cromatografiacutea liacutequida de altaresolucioacuten serie 983090983088983088 Perkin Elmer (Boston Ma USA)compuesto por un auto muestreador bomba cuaternariadegasificador detector de iacutendice de refraccioacuten y hornopara columnas El sistema se operoacute con el programacomputacional TotalChrom versioacuten 983094983090983089 Se usoacute una

columna Rezex RCM-monosaccharide Ca+983090

(983096 ) de 983091983088cm de longitud y 983095983096 mm de diaacutemetro Como fase moacutevilse empleoacute agua grado HPLC La velocidad de flujo fuede 983088983094 mLmiddotmin-983089 el volumen de muestra inyectado de983090983088 μL y el tiempo de corrida 983090983093 min La temperatura dela columna se mantuvo a 983096983093 ordmC Se elaboraron curvaspatroacuten usando estaacutendares comerciales de sacarosaglucosa y fructosa (Sigma MN) en concentraciones de983088983093 983089983088 983090983093 y 983093 mgmiddotmL -983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) En cada punto de las curvas se inyectoacute al menostres veces para obtener los datos del aacuterea asociada a

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The minimum ranged between 983095 and 983089983094 degC and eventhough in some periods it was less than 983089983088 degC it did notaffect plant growth and yield since it is considered thatpotato tolerates low temperatures with 983094 degC being itsbase temperature (Cao amp Tibbitts 983089983097983097983093)

The mean temperature was 983089983097 degC similar to thatreported by Cao and Tibbitts (983089983097983097983092) to obtain high

biomass and tuber production (983090983088 degC) Struik Haverkort Vreugdengil Bus and Dankert (983089983097983097983088) stress theimportance of high temperatures during RS 983089 as theypromote stolon branching an important condition for

yield because the number of tubers formed is closelyrelated to the number of stolons (Haverkort Van De

Waart amp Bodlaender 983089983097983097983088)

Crop Phenology

In general there were differential genotypic effects ( P le983088983088983093) in crop phenology The variety Alpha required moreGDD to reach the stolon (983089983094983095) tuberization (983090983094983089) and tuber

physiological maturity (983089983093983092) stages of the tuber its lifecycle was significantly longer than in the other varieties

Vivaldi was the variety that reached maturity the fastestbecause it needed 983089983093 fewer GDD than Alpha (Table 983091)

cada concentracioacuten con esta informacioacuten se obtuvieronlas ecuaciones de regresioacuten para cada azuacutecar

Con los datos obtenidos se realizaron un anaacutelisis de varianza y de regresioacuten ademaacutes se aplicaron pruebas decomparacioacuten de medias de Tukey para ello se empleoacuteel paquete Statistical Analysis System (SAS 983090983088983088983092)

Resultados y discusioacuten

Temperatura

La temperatura del aire fue fluctuante durante el ciclode cultivo (Figura 983089) la maacutexima varioacute entre 983089983095 y 983091983094 ordmCSuperiores a 983090983091 ordmC se registraron desde los 983096 hasta los983093983092 diacuteas despueacutes de la emergencia y coincidieron con laetapa vegetativa (EV) reproductiva inicial (ER 983089) y mitadde la reproductiva final (ER 983090) La miacutenima osciloacute entre 983095

y 983089983094 ordmC y a pesar de que en algunos periodos fue inferiora 983089983088 ordmC no afectoacute al crecimiento y rendimiento de laplanta ya que se considera que la papa resiste bajas

temperaturas siendo su base 983094 ordmC (Cao amp Tibbitts 983089983097983097983093)

La temperatura media fue de 983089983097 ordmC similar a lareportada por Cao y Tibbitts (983089983097983097983092) para obtener alta

Figure 983089 Air temperature recorded during the potato growing season under field conditions San Lorenzo

Cuapiaxtla Tlax Spring-Summer Cycle 983090983088983088983096 The arrows indicate the phases of emergence (E) stolon

initiation (SI) tuberization (T) and tuber physiological maturity (TM) and the intervals between them

at the vegetative (VS) initial reproductive (RS 983089) and final reproductive (RS 983090) stages Average values of

four varieties

Figura 983089 Temperatura del aire registrada durante el ciclo de cultivo de papa en condiciones de campo San

Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096 Las flechas indican las fases de emergencia

(E) inicio de estoloneo (ES) tuberizacioacuten (T) y madurez fisioloacutegica del tubeacuterculo (MT) y los intervalos

entre estas a las etapas vegetativa (EV) reproductiva inicial (ER 983089) y reproductiva final (ER 983090) Valores

promedio de cuatro variedades

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The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

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Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

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durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

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Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

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Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 518

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

at random from the experimental unit during apicaldominance (a single sprout) multiple sprouting (morethan one sprout) and senescence (more than onesprout and turgor loss in tubers) On each occasion the longest sprout was measured and from multiplesprouting all sprouts were measured for dry weight

Quantification of soluble carbohydrates by high-

performance liquid chromatography (HPLC)

Sample collection

Four samplings were performed during the storageperiod the first during dormancy and the other threeduring the physiological ages of apical dominancemultiple sprouting and senescence of the tuber ofeach variety On each occasion a tuber contained in thepaper bags was taken and a sample obtained from it

with two replications being made for the larger tubers Average tuber weights for the sampling process rangedbetween 983089983093983090 and 983089983096983089 g (Table 983090)

From the central part of each tuber between the basaland apical end a sample of 983093 plusmn 983089 g was extracted andplaced in liquid nitrogen (-983089983097983094 degC) until analysis

Extraction of soluble carbohydrates

The frozen tuber samples were individually ground ina blender with 983090983088 mL of 983095983088 HPLC-grade ethanol and

Condiciones de almacenamiento

En la cosecha los tubeacuterculos de cuatro plantas por unidadexperimental seleccionadas al azar fueron colocados enbolsas de papel para ser almacenados en laboratorio conluz ambiental bajo las condiciones de temperatura yhumedad relativa que se muestran en el Cuadro 983089 hastaque mostraron el maacuteximo desarrollo de brote

Se consideroacute que los tubeacuterculos contenidos en lasbolsas de papel dejaron la condicioacuten de reposo cuandoen 983093983088 de estos el brote principal midioacute 983092 plusmn 983090 mm Lalongitud (mm) y peso seco (mg) del brote se registroacuteen cinco tubeacuterculos elegidos al azar de la unidadexperimental en dominancia apical (un solo brote)brotacioacuten muacuteltiple (maacutes de un brote) y senescencia (maacutesde un brote y peacuterdida de turgencia de los tubeacuterculos)En cada ocasioacuten se midioacute el brote de mayor longitud ya partir de la brotacioacuten muacuteltiple se consideraron todoslos brotes para medir el peso seco

Cuantificacioacuten de carbohidratos solubles por croma-tografiacutea liacutequida de alta resolucioacuten (HPLC)

Obtencioacuten de la muestra

Durante el periodo de almacenamiento se realizaroncuatro muestreos el primero en condiciones dereposo y los otros tres correspondieron a las edadesfisioloacutegicas de dominancia apical brotacioacuten muacuteltiple

Table 983090 Seed-tuber weight at different physiological states of four potato varieties

Cuadro 983090 Peso de semilla-tubeacuterculo en diferentes estados fisioloacutegicos de cuatro variedades de papa

Seed-tuber hysiological state frasl Estado fisioloacutegico de semilla-tubeacuterculo

Variety frasl

VariedadDormancy frasl Reposo

Apical dominance frasl

Dominancia apical

Multiple sprouting frasl

Brotacioacuten muacuteltiple

Senescence frasl

Senescencia

DAHz frasl

DDCz

Weight y (g) frasl

Peso y (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

DAH frasl

DDC

Weight (g) frasl

Peso (g)

Alpha 983090983090 983089983097983089 983089983089983090 983089983096983090 983089983092983096 983089983094983091 983090983090983091 983089983093983088

Atlantic 983090983090 983089983092983090 983089983092983095 983089983093983089 983090983089983094 983089983092983093 983090983096983088 983089983091983088

Gigant 983090983090 983090983088983093 983095983095 983090983091983096 983097983096 983090983090983096 983090983090983091 983090983088983088

Vivaldi 983090983090 983089983096983096 983097983096 983089983092983094 983089983092983096 983089983092983097 983090983090983091 983089983091983088

Mean fraslMedia

983090983090 983089983096983089 983089983088983097 983089983095983097 983089983093983091 983089983095983089 983090983091983095 983089983093983090

zDDC days after harvest y average value of two tubers

zDDC diacuteas despueacutes de la cosecha y valor promedio de dos tubeacuterculos

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983091983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

brought to boiling temperature for 983093 min The extract was decanted and a second extraction was made fromthe residue with 983089983088 mL of ethanol at the end theextracts were mixed Subsequently the extract wascentrifuged for 983089983093 min at 983092983088983088983088 x g in a HERLE modelZ983090983091983088A centrifuge (Labnet International Inc Edison NJ

USA) filtered with Whatman No 983092 paper and mannitol was added at a concentration of 983092 mgmiddotmL-983089 as internal

standard Finally the extract was gauged to 983090983093 mL withthe same kind of solvent used for the extraction

Then 983089983088 mL of the extract were taken and concentratedto dryness with reduced pressure and low temperature(983092983088 degC) in a BUCHI R-983090983089983093 rotary evaporator (Switzerland)The residue was resuspended in 983090 mL of HPLC-grade

water in order to pass it through a column containingion-exchange resins one basic (983088983093 mL of Dowex - 983089 x983096 - Fluka) and the other acidic (983088983093 mL of Dowex - 983093983088W

X983096 - Fluka) at a ratio of 983089983089 (ww) The purified sample was passed through a 983088983092983093-μm nylon acrodisc in orderto collect the sample in a 983089983093 mL vial

HPLC sugar analysis

A Perkin Elmer series 983090983088983088 high-performance liquidchromatography system (Boston Ma USA) consistingof an autosampler quaternary pump degasserrefractive index detector and a column oven wasused The system is operated with TotalChrom version983094983090983089 software A Rezex RCM-monosaccharide Ca+983090 (983096 )column 983091983088 cm in length and 983095983096 mm in diameter wasused As the mobile phase HPLC-grade water was usedThe flow rate was 983088983094 mLmiddotmin-983089 the sample injection

volume 983090983088 μL and the running time 983090983093 min Columntemperature was maintained at 983096983093 degC Pattern curves

were prepared using commercial sucrose glucose andfructose standards (Sigma MN) at concentrations of983088983093 983089983088 983090983093 and 983093 mgmiddotmL-983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) At each point of the curves it was injected at leastthree times to obtain the data of the area associated

with each concentration area using this informationthe regression equations for each sugar were obtained

Analyses of variance and regression were performed with the data obtained Also Tukeyrsquos test was appliedusing the Statistical Analysis System package (SAS 983090983088983088983092)

Results and discussion

Temperature

The air temperature fluctuated during the growingseason (Figure 983089) with the maximum varying between983089983095 and 983091983094 degC Temperatures above 983090983091 degC were recordedfrom 983096 to 983093983092 days after emergence and coincided

with the vegetative (VS) and initial reproductive (RS983089)stages and half of the final reproductive stage (RS 983090)

y senescencia del tubeacuterculo de cada variedad En cadaocasioacuten se tomoacute un tubeacuterculo contenido en las bolsasde papel el de mayor tamantildeo en dos repeticiones conpesos promedio entre 983089983093983090 y 983089983096983089 g (Cuadro 983090) del que seobtuvo la muestra

De la parte central de cada tubeacuterculo entre el extremobasal y apical se extrajo una muestra de 983093 plusmn 983089 g que se

colocoacute en nitroacutegeno liacutequido (-983089983097983094 ordmC) hasta su anaacutelisis

Extraccioacuten de carbohidratos solubles

Las muestras congeladas de tubeacuterculo fueron molidasen licuadora individualmente con 983090983088 mL de etanolgrado HPLC a 983095983088 y llevadas a temperatura deebullicioacuten durante 983093 min El extracto se decantoacute yal residuo se le realizoacute una segunda extraccioacuten con983089983088 mL de etanol al final los extractos se mezclaronPosteriormente el extracto se centrifugoacute por 983089983093 mina 983092983088983088983088 x g en una centriacutefuga HERLE modelo Z983090983091983088A(Labnet International Inc Edison NJ USA) se filtroacute

con papel Whatman nuacutemero 983092 y se agregoacute manitol enconcentracioacuten de 983092 mgmiddotmL-983089 como estaacutendar internoFinalmente el extracto se aforoacute a 983090983093 mL con el mismotipo de solvente empleado para la extraccioacuten

Del extracto aforado se tomaron 983089983088 mL paraconcentrar a sequedad con presioacuten reducida y bajatemperatura (983092983088 ordmC) en un rotoevaporador BUCHIR-983090983089983093 (Switzerland) El residuo se resuspendioacute en 983090 mLde agua grado HPLC para pasarlo por una columnaque conteniacutea resinas intercambiadoras de iones unabaacutesica (983088983093 mL de Dowex - 983089 x 983096 - Fluka) y la otra aacutecida(983088983093 mL de Dowex - 983093983088W X983096 - Fluka) en proporcioacuten 983089983089(pp) La muestra purificada se pasoacute por un acrodiscode nylon de 983088983092983093 μm con el fin de colectar la muestraen un vial de 983089983093 mL

Anaacutelisis de azuacutecares en HPLC

Se utilizoacute un equipo de cromatografiacutea liacutequida de altaresolucioacuten serie 983090983088983088 Perkin Elmer (Boston Ma USA)compuesto por un auto muestreador bomba cuaternariadegasificador detector de iacutendice de refraccioacuten y hornopara columnas El sistema se operoacute con el programacomputacional TotalChrom versioacuten 983094983090983089 Se usoacute una

columna Rezex RCM-monosaccharide Ca+983090

(983096 ) de 983091983088cm de longitud y 983095983096 mm de diaacutemetro Como fase moacutevilse empleoacute agua grado HPLC La velocidad de flujo fuede 983088983094 mLmiddotmin-983089 el volumen de muestra inyectado de983090983088 μL y el tiempo de corrida 983090983093 min La temperatura dela columna se mantuvo a 983096983093 ordmC Se elaboraron curvaspatroacuten usando estaacutendares comerciales de sacarosaglucosa y fructosa (Sigma MN) en concentraciones de983088983093 983089983088 983090983093 y 983093 mgmiddotmL -983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) En cada punto de las curvas se inyectoacute al menostres veces para obtener los datos del aacuterea asociada a

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The minimum ranged between 983095 and 983089983094 degC and eventhough in some periods it was less than 983089983088 degC it did notaffect plant growth and yield since it is considered thatpotato tolerates low temperatures with 983094 degC being itsbase temperature (Cao amp Tibbitts 983089983097983097983093)

The mean temperature was 983089983097 degC similar to thatreported by Cao and Tibbitts (983089983097983097983092) to obtain high

biomass and tuber production (983090983088 degC) Struik Haverkort Vreugdengil Bus and Dankert (983089983097983097983088) stress theimportance of high temperatures during RS 983089 as theypromote stolon branching an important condition for

yield because the number of tubers formed is closelyrelated to the number of stolons (Haverkort Van De

Waart amp Bodlaender 983089983097983097983088)

Crop Phenology

In general there were differential genotypic effects ( P le983088983088983093) in crop phenology The variety Alpha required moreGDD to reach the stolon (983089983094983095) tuberization (983090983094983089) and tuber

physiological maturity (983089983093983092) stages of the tuber its lifecycle was significantly longer than in the other varieties

Vivaldi was the variety that reached maturity the fastestbecause it needed 983089983093 fewer GDD than Alpha (Table 983091)

cada concentracioacuten con esta informacioacuten se obtuvieronlas ecuaciones de regresioacuten para cada azuacutecar

Con los datos obtenidos se realizaron un anaacutelisis de varianza y de regresioacuten ademaacutes se aplicaron pruebas decomparacioacuten de medias de Tukey para ello se empleoacuteel paquete Statistical Analysis System (SAS 983090983088983088983092)

Resultados y discusioacuten

Temperatura

La temperatura del aire fue fluctuante durante el ciclode cultivo (Figura 983089) la maacutexima varioacute entre 983089983095 y 983091983094 ordmCSuperiores a 983090983091 ordmC se registraron desde los 983096 hasta los983093983092 diacuteas despueacutes de la emergencia y coincidieron con laetapa vegetativa (EV) reproductiva inicial (ER 983089) y mitadde la reproductiva final (ER 983090) La miacutenima osciloacute entre 983095

y 983089983094 ordmC y a pesar de que en algunos periodos fue inferiora 983089983088 ordmC no afectoacute al crecimiento y rendimiento de laplanta ya que se considera que la papa resiste bajas

temperaturas siendo su base 983094 ordmC (Cao amp Tibbitts 983089983097983097983093)

La temperatura media fue de 983089983097 ordmC similar a lareportada por Cao y Tibbitts (983089983097983097983092) para obtener alta

Figure 983089 Air temperature recorded during the potato growing season under field conditions San Lorenzo

Cuapiaxtla Tlax Spring-Summer Cycle 983090983088983088983096 The arrows indicate the phases of emergence (E) stolon

initiation (SI) tuberization (T) and tuber physiological maturity (TM) and the intervals between them

at the vegetative (VS) initial reproductive (RS 983089) and final reproductive (RS 983090) stages Average values of

four varieties

Figura 983089 Temperatura del aire registrada durante el ciclo de cultivo de papa en condiciones de campo San

Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096 Las flechas indican las fases de emergencia

(E) inicio de estoloneo (ES) tuberizacioacuten (T) y madurez fisioloacutegica del tubeacuterculo (MT) y los intervalos

entre estas a las etapas vegetativa (EV) reproductiva inicial (ER 983089) y reproductiva final (ER 983090) Valores

promedio de cuatro variedades

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The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

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Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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983089983091983096 Growth yield and sugar

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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983089983092983088 Growth yield and sugar

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

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983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

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Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

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Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 618

983089983091983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

brought to boiling temperature for 983093 min The extract was decanted and a second extraction was made fromthe residue with 983089983088 mL of ethanol at the end theextracts were mixed Subsequently the extract wascentrifuged for 983089983093 min at 983092983088983088983088 x g in a HERLE modelZ983090983091983088A centrifuge (Labnet International Inc Edison NJ

USA) filtered with Whatman No 983092 paper and mannitol was added at a concentration of 983092 mgmiddotmL-983089 as internal

standard Finally the extract was gauged to 983090983093 mL withthe same kind of solvent used for the extraction

Then 983089983088 mL of the extract were taken and concentratedto dryness with reduced pressure and low temperature(983092983088 degC) in a BUCHI R-983090983089983093 rotary evaporator (Switzerland)The residue was resuspended in 983090 mL of HPLC-grade

water in order to pass it through a column containingion-exchange resins one basic (983088983093 mL of Dowex - 983089 x983096 - Fluka) and the other acidic (983088983093 mL of Dowex - 983093983088W

X983096 - Fluka) at a ratio of 983089983089 (ww) The purified sample was passed through a 983088983092983093-μm nylon acrodisc in orderto collect the sample in a 983089983093 mL vial

HPLC sugar analysis

A Perkin Elmer series 983090983088983088 high-performance liquidchromatography system (Boston Ma USA) consistingof an autosampler quaternary pump degasserrefractive index detector and a column oven wasused The system is operated with TotalChrom version983094983090983089 software A Rezex RCM-monosaccharide Ca+983090 (983096 )column 983091983088 cm in length and 983095983096 mm in diameter wasused As the mobile phase HPLC-grade water was usedThe flow rate was 983088983094 mLmiddotmin-983089 the sample injection

volume 983090983088 μL and the running time 983090983093 min Columntemperature was maintained at 983096983093 degC Pattern curves

were prepared using commercial sucrose glucose andfructose standards (Sigma MN) at concentrations of983088983093 983089983088 983090983093 and 983093 mgmiddotmL-983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) At each point of the curves it was injected at leastthree times to obtain the data of the area associated

with each concentration area using this informationthe regression equations for each sugar were obtained

Analyses of variance and regression were performed with the data obtained Also Tukeyrsquos test was appliedusing the Statistical Analysis System package (SAS 983090983088983088983092)

Results and discussion

Temperature

The air temperature fluctuated during the growingseason (Figure 983089) with the maximum varying between983089983095 and 983091983094 degC Temperatures above 983090983091 degC were recordedfrom 983096 to 983093983092 days after emergence and coincided

with the vegetative (VS) and initial reproductive (RS983089)stages and half of the final reproductive stage (RS 983090)

y senescencia del tubeacuterculo de cada variedad En cadaocasioacuten se tomoacute un tubeacuterculo contenido en las bolsasde papel el de mayor tamantildeo en dos repeticiones conpesos promedio entre 983089983093983090 y 983089983096983089 g (Cuadro 983090) del que seobtuvo la muestra

De la parte central de cada tubeacuterculo entre el extremobasal y apical se extrajo una muestra de 983093 plusmn 983089 g que se

colocoacute en nitroacutegeno liacutequido (-983089983097983094 ordmC) hasta su anaacutelisis

Extraccioacuten de carbohidratos solubles

Las muestras congeladas de tubeacuterculo fueron molidasen licuadora individualmente con 983090983088 mL de etanolgrado HPLC a 983095983088 y llevadas a temperatura deebullicioacuten durante 983093 min El extracto se decantoacute yal residuo se le realizoacute una segunda extraccioacuten con983089983088 mL de etanol al final los extractos se mezclaronPosteriormente el extracto se centrifugoacute por 983089983093 mina 983092983088983088983088 x g en una centriacutefuga HERLE modelo Z983090983091983088A(Labnet International Inc Edison NJ USA) se filtroacute

con papel Whatman nuacutemero 983092 y se agregoacute manitol enconcentracioacuten de 983092 mgmiddotmL-983089 como estaacutendar internoFinalmente el extracto se aforoacute a 983090983093 mL con el mismotipo de solvente empleado para la extraccioacuten

Del extracto aforado se tomaron 983089983088 mL paraconcentrar a sequedad con presioacuten reducida y bajatemperatura (983092983088 ordmC) en un rotoevaporador BUCHIR-983090983089983093 (Switzerland) El residuo se resuspendioacute en 983090 mLde agua grado HPLC para pasarlo por una columnaque conteniacutea resinas intercambiadoras de iones unabaacutesica (983088983093 mL de Dowex - 983089 x 983096 - Fluka) y la otra aacutecida(983088983093 mL de Dowex - 983093983088W X983096 - Fluka) en proporcioacuten 983089983089(pp) La muestra purificada se pasoacute por un acrodiscode nylon de 983088983092983093 μm con el fin de colectar la muestraen un vial de 983089983093 mL

Anaacutelisis de azuacutecares en HPLC

Se utilizoacute un equipo de cromatografiacutea liacutequida de altaresolucioacuten serie 983090983088983088 Perkin Elmer (Boston Ma USA)compuesto por un auto muestreador bomba cuaternariadegasificador detector de iacutendice de refraccioacuten y hornopara columnas El sistema se operoacute con el programacomputacional TotalChrom versioacuten 983094983090983089 Se usoacute una

columna Rezex RCM-monosaccharide Ca+983090

(983096 ) de 983091983088cm de longitud y 983095983096 mm de diaacutemetro Como fase moacutevilse empleoacute agua grado HPLC La velocidad de flujo fuede 983088983094 mLmiddotmin-983089 el volumen de muestra inyectado de983090983088 μL y el tiempo de corrida 983090983093 min La temperatura dela columna se mantuvo a 983096983093 ordmC Se elaboraron curvaspatroacuten usando estaacutendares comerciales de sacarosaglucosa y fructosa (Sigma MN) en concentraciones de983088983093 983089983088 983090983093 y 983093 mgmiddotmL -983089 (Rodriacuteguez-Saona amp Wrolstad983089983097983097983095) En cada punto de las curvas se inyectoacute al menostres veces para obtener los datos del aacuterea asociada a

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

The minimum ranged between 983095 and 983089983094 degC and eventhough in some periods it was less than 983089983088 degC it did notaffect plant growth and yield since it is considered thatpotato tolerates low temperatures with 983094 degC being itsbase temperature (Cao amp Tibbitts 983089983097983097983093)

The mean temperature was 983089983097 degC similar to thatreported by Cao and Tibbitts (983089983097983097983092) to obtain high

biomass and tuber production (983090983088 degC) Struik Haverkort Vreugdengil Bus and Dankert (983089983097983097983088) stress theimportance of high temperatures during RS 983089 as theypromote stolon branching an important condition for

yield because the number of tubers formed is closelyrelated to the number of stolons (Haverkort Van De

Waart amp Bodlaender 983089983097983097983088)

Crop Phenology

In general there were differential genotypic effects ( P le983088983088983093) in crop phenology The variety Alpha required moreGDD to reach the stolon (983089983094983095) tuberization (983090983094983089) and tuber

physiological maturity (983089983093983092) stages of the tuber its lifecycle was significantly longer than in the other varieties

Vivaldi was the variety that reached maturity the fastestbecause it needed 983089983093 fewer GDD than Alpha (Table 983091)

cada concentracioacuten con esta informacioacuten se obtuvieronlas ecuaciones de regresioacuten para cada azuacutecar

Con los datos obtenidos se realizaron un anaacutelisis de varianza y de regresioacuten ademaacutes se aplicaron pruebas decomparacioacuten de medias de Tukey para ello se empleoacuteel paquete Statistical Analysis System (SAS 983090983088983088983092)

Resultados y discusioacuten

Temperatura

La temperatura del aire fue fluctuante durante el ciclode cultivo (Figura 983089) la maacutexima varioacute entre 983089983095 y 983091983094 ordmCSuperiores a 983090983091 ordmC se registraron desde los 983096 hasta los983093983092 diacuteas despueacutes de la emergencia y coincidieron con laetapa vegetativa (EV) reproductiva inicial (ER 983089) y mitadde la reproductiva final (ER 983090) La miacutenima osciloacute entre 983095

y 983089983094 ordmC y a pesar de que en algunos periodos fue inferiora 983089983088 ordmC no afectoacute al crecimiento y rendimiento de laplanta ya que se considera que la papa resiste bajas

temperaturas siendo su base 983094 ordmC (Cao amp Tibbitts 983089983097983097983093)

La temperatura media fue de 983089983097 ordmC similar a lareportada por Cao y Tibbitts (983089983097983097983092) para obtener alta

Figure 983089 Air temperature recorded during the potato growing season under field conditions San Lorenzo

Cuapiaxtla Tlax Spring-Summer Cycle 983090983088983088983096 The arrows indicate the phases of emergence (E) stolon

initiation (SI) tuberization (T) and tuber physiological maturity (TM) and the intervals between them

at the vegetative (VS) initial reproductive (RS 983089) and final reproductive (RS 983090) stages Average values of

four varieties

Figura 983089 Temperatura del aire registrada durante el ciclo de cultivo de papa en condiciones de campo San

Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096 Las flechas indican las fases de emergencia

(E) inicio de estoloneo (ES) tuberizacioacuten (T) y madurez fisioloacutegica del tubeacuterculo (MT) y los intervalos

entre estas a las etapas vegetativa (EV) reproductiva inicial (ER 983089) y reproductiva final (ER 983090) Valores

promedio de cuatro variedades

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983091983094 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

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durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

Ap Rees T amp Morrell S (983089983097983097983088) Carbohydrate metabolismin developing potatoes American Journal of Potato

Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

change patterns affect growth and tuberization inpotatoes Journal of the American Society for Horticultural

Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 718

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

The minimum ranged between 983095 and 983089983094 degC and eventhough in some periods it was less than 983089983088 degC it did notaffect plant growth and yield since it is considered thatpotato tolerates low temperatures with 983094 degC being itsbase temperature (Cao amp Tibbitts 983089983097983097983093)

The mean temperature was 983089983097 degC similar to thatreported by Cao and Tibbitts (983089983097983097983092) to obtain high

biomass and tuber production (983090983088 degC) Struik Haverkort Vreugdengil Bus and Dankert (983089983097983097983088) stress theimportance of high temperatures during RS 983089 as theypromote stolon branching an important condition for

yield because the number of tubers formed is closelyrelated to the number of stolons (Haverkort Van De

Waart amp Bodlaender 983089983097983097983088)

Crop Phenology

In general there were differential genotypic effects ( P le983088983088983093) in crop phenology The variety Alpha required moreGDD to reach the stolon (983089983094983095) tuberization (983090983094983089) and tuber

physiological maturity (983089983093983092) stages of the tuber its lifecycle was significantly longer than in the other varieties

Vivaldi was the variety that reached maturity the fastestbecause it needed 983089983093 fewer GDD than Alpha (Table 983091)

cada concentracioacuten con esta informacioacuten se obtuvieronlas ecuaciones de regresioacuten para cada azuacutecar

Con los datos obtenidos se realizaron un anaacutelisis de varianza y de regresioacuten ademaacutes se aplicaron pruebas decomparacioacuten de medias de Tukey para ello se empleoacuteel paquete Statistical Analysis System (SAS 983090983088983088983092)

Resultados y discusioacuten

Temperatura

La temperatura del aire fue fluctuante durante el ciclode cultivo (Figura 983089) la maacutexima varioacute entre 983089983095 y 983091983094 ordmCSuperiores a 983090983091 ordmC se registraron desde los 983096 hasta los983093983092 diacuteas despueacutes de la emergencia y coincidieron con laetapa vegetativa (EV) reproductiva inicial (ER 983089) y mitadde la reproductiva final (ER 983090) La miacutenima osciloacute entre 983095

y 983089983094 ordmC y a pesar de que en algunos periodos fue inferiora 983089983088 ordmC no afectoacute al crecimiento y rendimiento de laplanta ya que se considera que la papa resiste bajas

temperaturas siendo su base 983094 ordmC (Cao amp Tibbitts 983089983097983097983093)

La temperatura media fue de 983089983097 ordmC similar a lareportada por Cao y Tibbitts (983089983097983097983092) para obtener alta

Figure 983089 Air temperature recorded during the potato growing season under field conditions San Lorenzo

Cuapiaxtla Tlax Spring-Summer Cycle 983090983088983088983096 The arrows indicate the phases of emergence (E) stolon

initiation (SI) tuberization (T) and tuber physiological maturity (TM) and the intervals between them

at the vegetative (VS) initial reproductive (RS 983089) and final reproductive (RS 983090) stages Average values of

four varieties

Figura 983089 Temperatura del aire registrada durante el ciclo de cultivo de papa en condiciones de campo San

Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096 Las flechas indican las fases de emergencia

(E) inicio de estoloneo (ES) tuberizacioacuten (T) y madurez fisioloacutegica del tubeacuterculo (MT) y los intervalos

entre estas a las etapas vegetativa (EV) reproductiva inicial (ER 983089) y reproductiva final (ER 983090) Valores

promedio de cuatro variedades

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983091983094 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

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Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

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Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

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Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

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Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

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Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 818

983089983091983094 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

The vegetative and initial reproductive stagesaccounted for 983092983093 of the total duration of the cropcycle in Alpha unlike the other varieties in which theyaccounted for only 983090983094 while the final reproductivestage accounted for 983093983093 in Alpha and 983095983092 in the other

varieties (Table 983091) The greater precocity of Vivaldi with respect to the other varieties may be due to thelower GDD requirement in the vegetative and initial

reproductive stages because the shorter duration ofthese stages affects the crop growth period (KoomanFahem Tegera amp Haverkort 983089983097983097983094)

Yield and its components

Yield expressed as total tuber weight per plant wassimilar ( P le 983088983088983093) in the varieties Alpha Gigant and

Vivaldi (983094983093983097 g) which had a 983093983096 greater yield than the variety Atlantic (Table 983092)

The components that contributed most to the yieldsof the varieties were the number and weight of

commercial and non-commercial tubers per plant inthe varieties Alpha and Vivaldi and number weightof commercial tubers per plant and average tuber

weight per plant in Gigant The variety Alpha had asignificantly higher total number of tubers per plantexceeding Atlantic by 983094983088 and Gigant and Vivaldi by983090983097 (Table 983092) however it had a lower proportion ofcommercial tubers per plant (983094983095 ) compared to theother varieties (983095983090 ) This indicates that the increase in

produccioacuten de biomasa y tubeacuterculos (983090983088 ordmC) StruikHaverkort Vreugdengil Bus y Dankert (983089983097983097983088) destacanla importancia de la presencia de altas temperaturasdurante la ER 983089 ya que eacutestas promueven la ramificacioacutendel estoloacuten condicioacuten importante para el rendimientopuesto que el nuacutemero de tubeacuterculos formados estaacuteestrechamente relacionado con el nuacutemero de estolones(Haverkort Van De Waart amp Bodlaender 983089983097983097983088)

Fenologiacutea del cultivo

En general hubo efectos genotiacutepicos diferenciales( P le 983088983088983093) en la fenologiacutea del cultivo La variedad

Alpha requirioacute maacutes GD para alcanzar las fasesde estoloneo (983089983094983095) tuberizacioacuten (983090983094983089) y madurezfisioloacutegica del tubeacuterculo (983089983093983092) su ciclo bioloacutegico fuesignificativamente maacutes prolongado que en las demaacutes

variedades Vivaldi fue la variedad que alcanzoacute maacutesraacutepidamente la madurez pues necesitoacute 983089983093 menos GDque Alpha (Cuadro 983091)

La etapa vegetativa y reproductiva inicial contribuyeroncon 983092983093 en la duracioacuten total del ciclo de cultivo en

Alpha a diferencia de las otras variedades en quesolo fue 983090983094 en tanto que la etapa reproductivafinal contribuyoacute con 983093983093 en Alpha y con 983095983092 en las

variedades (Cuadro 983091) La mayor precocidad de Vivaldicon respecto al resto de las variedades se puede deberal menor requerimiento de GD en la etapa vegetativa

y reproductiva inicial ya que la menor duracioacuten de

Table 983091 Growing degree-day requirements to reach the main phenological phases and stages of four potato

varieties grown under field conditions San Lorenzo Cuapiaxtla Tlax Spring-Summer 983090983088983088983096

Cuadro 983091 Requerimientos de grados diacutea para alcanzar las principales fases y etapas fenoloacutegicas de cuatro

variedades de papa bajo condiciones de campo San Lorenzo Cuapiaxtla Tlax Ciclo Primavera-Verano

983090983088983088983096

Phases (GDD) frasl Fases (GD) STAGES (GDD) frasl Etapas (GD)

Variety frasl Variedad SI frasl ES T TM frasl MT VS frasl EV RS 983089 frasl ER 983089 RS 983090 frasl ER 983090

Alpha 983091983091983089az 983093983090983092a 983089983089983095983088a 983091983091983089a 983089983097983091a 983094983092983094c

Atlantic 983089983095983096b 983090983095983092c 983089983088983088983093bc 983089983095983096b 983097983093b 983095983091983089b

Gigant 983089983097983094b 983091983088983091b 983089983088983093983090b 983089983097983094b 983089983088983095b 983095983092983097ab

Vivaldi 983089983089983095c 983090983089983091d 983097983097983088c 983089983089983095c 983097983094b 983095983095983094a

Mean frasl Media 983090983088983093 983091983090983096 983089983088983093983092 983090983088983093 983089983090983091 983095983090983093

HSD frasl DMSH 983091983091 983090983091 983093983095 983091983091 983089983096 983091983096

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant differenceGDD growing degree-days SI stolon initiation T tuber initiation TM tuber physiological maturity VS vegetative stage RS 1 initial reproductive stage RS 2final reproductive stage

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaGD grados diacutea ES inicio de estoloneo T inicio de tuberizacioacuten MT madurez fisioloacutegica del tubeacuterculo EV etapa vegetativa ER 1 etapa reproductiva inicial ER2 etapa reproductiva final

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Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

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Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

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Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

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acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

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Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

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Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

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Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 918

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Table 983092 Yield and its components of four potato varieties grown under field conditions San Lorenzo Cuapiaxtla

Tlax Spring-Summer cycle 983090983088983088983096

Cuadro 983092 Rendimiento y sus componentes de cuatro variedades de papa bajo condiciones de campo San Lorenzo

Cuapiaxtla Tlax Ciclo Primavera-Verano 983090983088983088983096

Variety frasl

VariedadNCTP frasl NTCP

WCTP (g) frasl

PTCP (g)

NNCTP frasl

NTNCP

WNCTP (g) frasl

PTNCP (g)

TNTP frasl

NTTP

TWTP (g) frasl

PTTP (g)

ATWP (g)

frasl PPTP (g)

Alpha 983095983093az 983094983089983097a 983091983095a 983096983094a 983089983089983090a 983095983088983093a 983094983091b

Atlantic 983091983093b 983090983093983093b 983089983088b 983090983091b 983092983093c 983090983095983096b 983094983090b

Gigant 983094983088ab 983094983094983094a 983089983094b 983091983094b 983095983094b 983095983088983090a 983097983090a

Vivaldi 983093983090ab 983092983097983088a 983091983088a 983096983089a 983096983090b 983093983095983089a 983095983088ab

Mean frasl Media 983093983093 983093983088983095 983090983091 983093983094 983095983097 983093983094983092 983095983090

HSD frasl DMSH 983090983096 983090983089983097 983089983090 983091983097 983090983096 983090983088983090 983090983093

z Values with the same letter within columns are not statistically different (Tukey at P le 005)

HSD Honestly significant differenceNCTP number of commercial tubers per plant WCTP weight of commercial tubers per plant NNCTP Number of non-commercial tubers per plant WNCTP

weight of non-commercial tubers per plant TNTP total number of tubers per plant TWTP total weight of tubers per plant ATWP average tuber weight perplant

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaNTCP nuacutemero de tubeacuterculos comerciales por planta PTCP peso de tubeacuterculos comerciales por planta NTNCP nuacutemero de tubeacuterculos no comerciales por plantaPTNCP peso de tubeacuterculos no comerciales por planta NTTP nuacutemero total de tubeacuterculos por planta PTTP peso total de tubeacuterculos por planta PPTP pesopromedio de tubeacuterculo por planta

the number of tubers per plant might affect their finalsize (Walworth amp Carling 983090983088983088983090)

Sugar content in potato tubers

In general the concentration of soluble sugars in tubers was higher during sprouting than dormancy whenthey had on average 983091983093 983091983097 983091983095 and 983091983095 lower sucroseglucose fructose and total sugar contents respectivelythan in tubers with a different physiological age (Table983093) These results agree with those reported by SinhaCash and Chase (983089983097983097983090) and Sabba et al (983090983088983088983095) who statethat the least amount of sugars occurs during tuberdormancy a situation that can be attributed initially tofoliage senescence before harvest and consequently areduction in the transport of sugars and the conversion

of them into starch in the tuber (Stark amp Love 983090983088983088983091)

As tubers left dormancy their sugar content increased( P le 983088983088983093) Similar behavior was observed from apicaldominance to tuber senescence (Table 983093) In this regardSowokinos (983090983088983088983089) indicates that after tubers have beenstored for prolonged periods they naturally experienceincreases in sugar content due to the conversionof starch into glucose and fructose These changesare most evident when tubers are subjected to lowtemperatures as occurred in November December and

January in the present study

las etapas afecta el periodo de crecimiento del cultivo(Kooman Fahem Tegera amp Haverkort 983089983097983097983094)

Rendimiento y sus componentes

El rendimiento expresado en peso total de tubeacuterculospor planta fue similar ( P le 983088983088983093) en las variedades AlphaGigant y Vivaldi (983094983093983097 g) que presentaron rendimiento983093983096 mayor que Atlantic (Cuadro 983092)

Los componentes que maacutes contribuyeron sobre losrendimientos de las variedades fueron el nuacutemero y peso detubeacuterculos comerciales y no comerciales por planta en las

variedades Alpha y Vivaldi y nuacutemero peso de tubeacuterculoscomerciales por planta y peso promedio de tubeacuterculo porplanta enGigant La variedad Alpha tuvo significativamente

mayor nuacutemero total de tubeacuterculos por planta al superar983094983088 a Atlantic y en 983090983097 a Gigant y Vivaldi (Cuadro 983092)no obstante presentoacute menor proporcioacuten de tubeacuterculoscomerciales por planta (983094983095 ) con respecto a las otras

variedades (983095983090 ) Lo anterior indica que el incrementoen el nuacutemero de tubeacuterculos por planta podriacutea afectar sutamantildeo final (Walworth amp Carling 983090983088983088983090)

Contenido de azuacutecares en tubeacuterculos de papa

En general la concentracioacuten de azuacutecares solubles enel tubeacuterculo fue mayor durante la brotacioacuten que en la

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Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

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durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

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Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

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significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

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Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

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Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

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Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

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acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1018

983089983091983096 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Table 983093 Sugar content in tubers of four potato varieties at different physiological states under storage conditions

Chapingo Mex 983090983088983088983096-983090983088983088983097

Cuadro 983093 Contenido de azuacutecares en tubeacuterculos de cuatro variedades de papa en diferentes estados fisioloacutegicos

bajo condiciones de almacenamiento Chapingo Meacutex 983090983088983088983096-983090983088983088983097

Physiological state frasl

Bestado fisioloacutegico

Sugar concentration (mgmiddotg-983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl Sacarosa Glucose fraslGlucosa Fructose fraslFructosa Total frasl Totales

Reposo frasl Dormancy 983090983089983090bz 983089983094983095b 983088983097983091b 983092983095983091b

Dominancia apical frasl Apical dominance

983091983092983089a 983090983091983096ab 983089983091983089ab 983095983089983088a

Brotacioacuten muacuteltiple fraslMultiple sprouting

983091983088983097a 983090983097983089a 983089983093983090a 983095983093983091a

Senescencia frasl Senescence 983091983090983096a 983091983088983089a 983089983093983097a 983095983096983097a

HSD frasl DMSH 983088983095983096 983088983095983092 983088983093983096 983089983097983091

z Values with the same letter within columns are not statistically different (Tukey at P le 005)HSD honestly significant difference

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honesta

By analyzing the soluble sugars at dormancy and bytuber physiological age (Figure 983090) it was found thatthe total sugar content was similar among varieties( P le 983088983088983093) However the varieties Vivaldi and Giganthad a higher concentration of sucrose and glucose(983090983096 and 983090983097 respectively) whereas Alpha and Gigant

were 983092983091 higher in f ructose (Figure 983090a)

During apical dominance (Figure 983090b) multiple sprouting(Figure 983090c) and senescence (Figure 983090d) of the tubers the

variety Vivaldi recorded the highest ( P le 983088983088983093) amountof glucose fructose and total sugars moreover in theapical dominance condition it had 983092983089 higher sucrosecontent than the other varieties studied

The interaction between varieties and tuberphysiological age indicated that in soluble sugarcontent Vivaldi had significantly greater variationthrough the physiological states (Table 983094) In thisrespect studies by Sowokinos (983090983088983088983089) indicate that the

storage conditions and genetic load of the varietiesmay affect the sugar content in the tuber as there are varieties that are more susceptible to the influence ofenvironmental factors (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

The variety Alpha at dormancy had 983094983095 less sucrosethan Vivaldi had in apical dominance whereas the

varieties Alpha Atlantic and Vivaldi recorded 983094983097 and983093983096 less glucose and total sugars than Vivaldi inthe other sprouting states The variety Atlantic at

condicioacuten de reposo ya que en promedio tuvieron 983091983093983091983097 983091983095 y 983091983095 menor contenido de sacarosa glucosafructosa y azuacutecares totales respectivamente que enaquellos tubeacuterculos con diferente edad fisioloacutegica(Cuadro 983093) Estos resultados concuerdan con losreportados por Sinha Cash y Chase (983089983097983097983090) y Sabbaet al (983090983088983088983095) quienes indican que la menor cantidadde azuacutecares se presentan durante el reposo delos tubeacuterculos situacioacuten que puede ser atribuidainicialmente a la senescencia del follaje previo a lacosecha y en consecuencia reduccioacuten en el transportede azuacutecares y conversioacuten de estos a almidoacuten en eltubeacuterculo (Stark amp Love 983090983088983088983091)

A medida que los tubeacuterculos dejaron el estado dereposo aumentoacute su contenido de azuacutecares ( P le 983088983088983093)un comportamiento similar se observoacute a partir de ladominancia apical y hasta la senescencia del tubeacuterculo(Cuadro 983093) Al respecto Sowokinos (983090983088983088983089) indica quedespueacutes de que los tubeacuterculos son almacenados por

periodos prolongados experimentan de manera naturalincrementos en el contenido de azuacutecares debido a laconversioacuten de almidoacuten a glucosa y fructosa Estoscambios son maacutes evidentes cuando los tubeacuterculos sonsometidos a bajas temperaturas como ocurrioacute en losmeses de noviembre diciembre y enero en la presenteinvestigacioacuten

Al analizar los azuacutecares solubles en condicioacuten de reposo y por edad fisioloacutegica del tubeacuterculo (Figura 983090) se observoacuteque el contenido de azuacutecares totales fue similar entre

variedades ( P le 983088983088983093) Sin embargo las variedades Vivaldi

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1118

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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983089983092983088 Growth yield and sugar

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

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Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

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de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

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Figure 983090 Soluble sugar content in potato tubers in the states of a) dormancy (983090983090 DAH) b) apical dominance

(983089983088983097 DAH) c) multiple sprouting (983089983093983091 DAH) and d) senescence (983090983091983095 DAH) under storage conditions

Chapingo Mex Values with the same letter within each type of sugar are not statistically different

(Tukey P le 983088983088983093)

Figura 983090 Contenido de azuacutecares solubles en tubeacuterculos de papa en los estados de a) reposo (983090983090 DDC) b)

dominancia apical (983089983088983097 DDC) c) brotacioacuten muacuteltiple (153 DDC) y d) senescencia (237 DDC) en condiciones

de almacenamiento Chapingo Meacutex Valores con la misma letra dentro de cada tipo de azuacutecar no

difieren estadiacutesticamente (Tukey P le 983088983088983093)

y Gigant tuvieron mayor concentracioacuten de sacarosa yglucosa (983090983096 y 983090983097 respectivamente) mientras que Alpha

y Gigant fueron 983092983091 superiores en fructosa (Figura 983090a)

Durante la dominancia apical (Figura 983090b) brotacioacutenmuacuteltiple (Figura 2c) y senescencia (Figura 2d) de lostubeacuterculos la variedad Vivaldi registroacute la mayor( P le 983088983088983093) cantidad de glucosa fructosa y azuacutecarestotales ademaacutes en la condicioacuten de dominanciaapical tuvo 983092983089 mayor contenido de sacarosa que elresto de variedades estudiadas

La interaccioacuten entre variedades y edad fisioloacutegicadel tubeacuterculo indicoacute que el contenido de azuacutecaressolubles en Vivaldi presentoacute significativamente mayor

variacioacuten a traveacutes de los estados fisioloacutegicos (Cuadro983094) Al respecto estudios hechos por Sowokinos (983090983088983088983089)indican que las condiciones de almacenamiento yla carga geneacutetica de las variedades pueden afectar elcontenido de azuacutecares en el tubeacuterculo ya que existen

variedades que son maacutes susceptibles a la influenciade factores ambientales (Park et al 983090983088983088983097 Pritchard amp

Adam 983089983097983097983090)

dormancy was 983095983095 lower in fructose than Vivaldi inthe senescence state (Table 983094) These results coincide

with those of Sinha et al (983089983097983097983090) who obtained theleast amount of sugars during tuber dormancy Thismay be due to the ability of the genotypes to convertsoluble sugars into starch (Stark amp Love 983090983088983088983091) coupled

with decreased transport of sugars to the tuber due tosenescence or foliage defoliation

The variety Vivaldi recorded the highest sucrose (DA)glucose (BM and S) fructose (S) and total sugars (DA

to S) contents among all varieties and physiologicalstates (Table 983094) Sowokinos (983090983088983088983089) indicates thattubers after dormancy experience an increase in sugarcontent due to the breakdown of starch into glucoseand fructose therefore the tuber becomes a sourceof energy and substrates for subsequent processes(Vreugdenhil 983090983088983088983092)

Tuber sprouting

The appearance of the tuber sprout marks the end ofdormancy (Suttle 983090983088983088983092) From apical dominance until

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Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

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due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

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durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

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Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

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Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

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Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

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Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

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Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1218

983089983092983088 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Table 983094 Comparison of means of variety interaction by physiological state with the sugar content of potato

tubers under storage conditions Chapingo Mex

Cuadro 983094 Comparacioacuten de medias de la interaccioacuten variedad por estado fisioloacutegico del contenido de azuacutecares de

tubeacuterculos de papa en condiciones de almacenamiento Chapingo Meacutex

Variety frasl

Variedad

Physiological state frasl

Estado fisioloacutegico

Sugar concentration (mgmiddotg -983089 from sample expressed in fresh weight) frasl

Concentracioacuten de azuacutecares (mgmiddotg-983089 de muestra expresado en peso fresco)

Sucrose frasl

Sacarosa

Glucose frasl

Glucosa

Fructose frasl

Fructosa Total frasl Totales

Alpha D frasl R (983090983090 DDC) 983089983094983089dz 983089983093983094d 983089983089983090de 983092983091983088c

Alpha AD frasl DA (983089983089983090 DDC) 983089983096983089cd 983090983089983095bcd 983089983089983091de 983093983089983089bc

Alpha MS frasl BM (983089983092983096 DDC) 983090983092983089bcd 983090983090983088bcd 983089983091983090cde 983093983097983091bc

Alpha S (983090983090983091 DDC) 983091983088983095bcd 983090983090983091bcd 983089983089983093de 983094983092983093bc

Atlantic D frasl R (983090983090 DDC) 983089983097983096cd 983089983091983097d 983088983094983092e 983092983088983091c

Atlantic AD frasl DA (983089983092983095 DDC) 983091983095983095ab 983089983092983093d 983088983097983090de 983094983089983092bc

Atlantic MS frasl BM (983090983089983094 DDC) 983091983095983094ab 983089983096983095cd 983088983097983091de 983094983093983095bc

Atlantic S (983090983096983088 DDC) 983091983091983095abc 983089983096983093cd 983088983096983097de 983094983089983089bc

Gigant D frasl R (983090983090 DDC) 983090983090983090bcd 983090983089983091bcd 983089983090983092cde 983093983093983097bc

Gigant AD frasl DA (983095983095 DDC) 983091983089983091bcd 983090983091983091bcd 983089983089983089de 983094983093983095bc

Gigant MS frasl BM (983097983096 DDC) 983090983097983091bcd 983090983094983095bcd 983089983093983090bcde 983095983089983090abc

Gigant S (983090983090983091 DDC) 983091983093983093abc 983091983089983093bc 983089983093983095bcd 983096983090983095ab

Vivaldi D frasl R (983090983090 DDC) 983090983094983097bcd 983089983094983088d 983088983095983089de 983093983088983089bc

Vivaldi AD frasl DA (983097983096 DDC) 983092983097983091a 983091983093983094ab 983090983089983088abc 983089983088983093983097a

Vivaldi MS frasl BM (983089983092983096 DDC) 983091983090983096abcd 983092983097983090a 983090983091983089ab 983089983088983093983090a

Vivaldi S (983090983090983091 DDC) 983091983089983092bcd 983092983096983090a 983090983095983095a 983089983088983095983091a

HSD frasl DMSH 983089983095983092 983089983092983095 983088983097983088 983091983096983094

z Values with the same letter within columns are not significantly different (Tukey at P le 005)HSD honestly significant differenceD dormancy AD apical dominance MS multiple sprouting S senescence DAH days after harvest

z Valores con la misma letra dentro de columnas no difieren estadiacutesticamente (Tukey a una P le 005)DMSH diferencia miacutenima significativa honestaR reposo DA dominancia apical BM brotacioacuten muacuteltiple S senescencia DDC diacuteas despueacutes de la cosecha

La variedad Alpha en condiciones de reposo tuvo983094983095 menos sacarosa que Vivaldi en la dominanciaapical en tanto que las variedades Alpha Atlantic

y Vivaldi registraron 983094983097 y 983093983096 menor cantidad deglucosa y azuacutecares totales que Vivaldi en los demaacutesestados de brotacioacuten La variedad Atlantic en reposofue 983095983095 inferior en fructosa que Vivaldi en el estadode senescencia (Cuadro 983094) Estos resultados concuerdancon los de Sinha et al (983089983097983097983090) quienes obtuvieron lamenor cantidad de azuacutecares durante el reposo delos tubeacuterculos de papa Lo anterior puede ser debidoa la habilidad de los genotipos para la conversioacuten de

multiple sprouting of the tuber the v arieties Gigantand Vivaldi had significantly greater ( P le 983088983088983093) sproutlength (983091983089 and 983091983092 ) (Figure 983091a) and dry weight (983094983094and 983093983097 ) (Figure 983091b) respectively than the varieties

Atlantic and Alpha In senescent tubers sprout lengthand dry weight reached the highest values The

varieties Gigant Vivaldi and Atlantic had 983091983092 longersprouts than Alpha Gigant had 983094983088 higher sprout dry

weight than Alpha and Atlantic (Figure 983091)

Some research indicates that the diversification among varieties in sprout length and dry matter could be

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1318

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1418

983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

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Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

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Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

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Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1318

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

due to the length of the crop cycle because duringtuber sprouting sprout length is shorter in late thanearly varieties (Knowles Driskill amp Knowles 983090983088983088983097) asoccurred with the Alpha variety in the present study

which had the longest life cycle Other studies attributeit to tuber size (Park et al 983090983088983088983097) and the ability varietieshave to breakdown the starch into reducing sugars

which are remobilized towards sprout formation and

growth (Vreugdenhil 983090983088983088983092)

azuacutecares solubles hacia almidoacuten (Stark amp Love 983090983088983088983091)aunado a la disminucioacuten del transporte de azuacutecares altubeacuterculo por la senescencia o defoliacioacuten del follajeLa variedad Vivaldi registroacute el mayor contenido desacarosa (DA) glucosa (BM y S) fructosa (S) y azuacutecarestotales (DA a S) entre todas las variedades y estadosfisioloacutegicos (Cuadro 983094) Sowokinos (983090983088983088983089) indica quelos tubeacuterculos despueacutes del reposo experimentan un

incremento en el contenido de azuacutecares debido al

Figure 983091 Sprout length (a) and dry weight (b) in the physiological states of apical dominance (AD) multiple

sprouting (MS) and senescence (S) in four potato varieties under storage conditions Chapingo Mex

Values with the same letter within each state are not statistically different (Tukey P le 983088983088983093)

Figura 983091 Longitud (a) y peso seco (b) de brote en los estados fisioloacutegicos de dominancia apical (DA) brotacioacuten

muacuteltiple (BM) y senescencia (S) en cuatro variedades de papa bajo condiciones de almacenamiento

Chapingo Meacutex Valores con la misma letra dentro de cada estado no difieren estadiacutesticamente (Tukey

P le 983088983088983093)

Sugar content in the tuber sprouting process

During tuber sprouting sprout length and dry weighthad a high significant association with soluble sugarcontent (Figure 983092) Increased sprout length (Figure 983092aand 983092b) and weight (Figure 983092c and 983092d) were accompaniedby an increase on average in the concentration ofreducing (glucose and fructose) and total sugars inapical dominance multiple sprouting and senescenceof the four varieties

In the varieties Alpha (Figure 983093a and 983093b) and Gigant(Figure 983093c and 983093d) sprout length and weight increasedas total sugars did Vivaldi showed similar behavior inthe content of reducing sugars (Figure 983093e and 983093f)

Sugar content at harvest is one of the most importantparameters that determine maturity and duringsprouting sprout vigor in potato tubers (Ap Reesamp Morrell 983089983097983097983088) In the present study the averagebehavior of varieties (Figure 983092) and individuallyin Alpha Gigant and Vivaldi (Figure 983093) showed

desdoblamiento del almidoacuten a glucosa y fructosa por loque el tubeacuterculo se convierte en una fuente que proveeenergiacutea y sustratos para los procesos subsecuentes(Vreugdenhil 983090983088983088983092)

Brotacioacuten de los tubeacuterculos

La aparicioacuten del brote en el tubeacuterculo marca el fin delestado de reposo (Suttle 983090983088983088983092) A partir de la dominanciaapical y hasta la brotacioacuten muacuteltiple del tubeacuterculo las

variedades Gigant y Vivaldi tuvieron significativamente

mayor ( P le 983088983088983093) longitud (983091983089 y 983091983092 ) (Figura 983091a) y pesoseco del brote (983094983094 y 983093983097 ) (Figura 983091b) respectivamenteal de Atlantic y Alpha En los tubeacuterculos senescentes lalongitud y peso seco del brote alcanzaron los mayores

valores Las variedades Gigant Vivaldi y Atlantic tuvieronbrotes 983091983092 maacutes largos que Alpha Gigant tuvo 983094983088 mayor peso seco de brote que Alpha y Atlantic (Figura 983091)

Algunas investigaciones indican que la diversificacioacutenentre variedades en longitud y materia seca del brotepodriacutea deberse a la duracioacuten del ciclo de cultivo ya que

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1418

983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1518

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1618

983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

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Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

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Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

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physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1418

983089983092983090 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

durante la brotacioacuten del tubeacuterculo la longitud del brotees maacutes corta en variedades tardiacuteas que en las precoces(Knowles Driskill amp Knowles 983090983088983088983097) como ocurrioacute conla variedad Alpha en la presente investigacioacuten que fue la

de mayor ciclo bioloacutegico Otros estudios lo atribuyen altamantildeo del tubeacuterculo (Park et al 983090983088983088983097) y a la habilidadque tienen las variedades para desdoblar el almidoacutena azuacutecares reductores su removilizacioacuten hacia laformacioacuten y crecimiento del brote (Vreugdenhil 983090983088983088983092)

Contenido de azuacutecares en el proceso de brotacioacutendel tubeacuterculo

Durante la brotacioacuten de los tubeacuterculos la longitud y peso seco del brote tuvieron alta asociacioacuten

increased sprout length and dry weight during apicaldominance multiple sprouting and senescence as thecontent of reducing and total sugars increased Theseresults agree with those reported by Davies (983089983097983097983088)

and Dogras et al (983089983097983096983097) who during tuber sproutingrecorded high reducing and total sugar contentspossibly due to the ability of some varieties to breakdown starch into glucose (Vreugdenhil 983090983088983088983092) to beused as a source of energy and substrates for sproutdevelopment Other studies indicate that when thepotato tuber reaches maturity the capacity for starchsynthesis decreases while the accumulation of sugarsin the tuber increases which contributes to thesprouting process (Viola et al 983090983088983088983095) as occurred inthis work

Figure 983092 Relationship between sprout length (a and b) and dry weight (c and d) with the content of reducing and

total sugars in four potato varieties under storage conditions Chapingo Mex The three points in each

graph represent the states of apical dominance (983089983088983097 DAH) multiple sprouting (983089983093983091 DAH) and senescence

(983090983091983095 DAH)

Figura 983092 Relacioacuten entre la longitud (a y b) y peso seco (c y d) del brote y el contenido de azuacutecares reductores y

totales en cuatro variedades de papa bajo condiciones de almacenamiento Chapingo Meacutex Los tres

puntos en cada graacutefica representan a los estados de dominancia apical (983089983088983097 DDC) brotacioacuten muacuteltiple

(153 DDC) y senescencia (237 DDC)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1518

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1618

983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

Ap Rees T amp Morrell S (983089983097983097983088) Carbohydrate metabolismin developing potatoes American Journal of Potato

Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

change patterns affect growth and tuberization inpotatoes Journal of the American Society for Horticultural

Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1718

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1518

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

Figure 983093 Relationship between sprout length and dry weight with total sugar content in the varieties Alpha (a

and b) and Gigant (c and d) and with reducing sugars in Vivaldi (e and f) under storage conditions

Chapingo Mex The three points in each graph represent the states of apical dominance (AD) multiple

sprouting (MS) and senescence (S)

Figura 983093 Relacioacuten entre la longitud y peso seco del brote con el contenido de azuacutecares totales en las variedades

Alpha (a y b) y Gigant (c y d) y con los azuacutecares reductores en Vivaldi (e y f) bajo condiciones de

almacenamiento Chapingo Meacutex Los tres puntos en cada graacutefica representan a los estados de

dominancia apical (DA) brotacioacuten muacuteltiple (BM) y senescencia (S)

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1618

983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

Ap Rees T amp Morrell S (983089983097983097983088) Carbohydrate metabolismin developing potatoes American Journal of Potato

Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

change patterns affect growth and tuberization inpotatoes Journal of the American Society for Horticultural

Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1718

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1618

983089983092983092 Growth yield and sugar

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

significativa con el contenido de azuacutecares solubles(Figura 983092) El incremento en la longitud (Figura 983092a

y 983092b) y peso del brote (Figura 983092c y 983092d) estuvieronacompantildeados del aumento en la concentracioacuten deazuacutecares reductores (glucosa y f ructosa) y totales en ladominancia apical brotacioacuten muacuteltiple y senescenciaen promedio de las cuatro variedades

En las variedades Alpha (Figura 983093a y 983093b) y Gigant(Figura 983093c y 983093d) aumentoacute la longitud y peso seco delbrote conforme los azuacutecares totales lo hicieron similarcomportamiento mostroacute Vivaldi en el contenido deazuacutecares reductores (Figura 983093e y 983093f)

El contenido de azuacutecares al momento de la cosecha esuno de los paraacutemetros maacutes importantes que determinanla madurez y durante la brotacioacuten el vigor del brote en lostubeacuterculos de papa (Ap Rees amp Morrell 983089983097983097983088) En la presenteinvestigacioacuten el comportamiento promedio de variedades(Figura 983092) y de manera individual en Alpha Gigant y

Vivaldi (Figura 983093) mostraron incremento en la longitud

y peso seco del brote en la dominancia apical brotacioacutenmuacuteltiple y senescencia a medida que el contenido deazuacutecares reductores y totales aumentoacute Estos resultadosconcuerdan con los reportados por Davies (983089983097983097983088) y Dograset al (983089983097983096983097) quienes durante la brotacioacuten de los tubeacuterculosregistraron alto contenido de azuacutecares reductores ytotales debido posiblemente a la habilidad que tienenalgunas variedades para desdoblar el almidoacuten a glucosa(Vreugdenhil 983090983088983088983092) y ser utilizada como fuente de energiacutea

y sustratos para el desarrollo del brote Otros estudiosindican que al momento de la madurez del tubeacuterculode papa la capacidad de siacutentesis de almidoacuten disminuyemientras que la acumulacioacuten de azuacutecares en el tubeacuterculoaumenta lo que contribuye al proceso de brotacioacuten (Violaet al 983090983088983088983095) como ocurrioacute en este trabajo

Ademaacutes de los azuacutecares (Viola et al 983090983088983088983095) y las hormonas(Sonnewald amp Sonnewald 983090983088983089983092) el genotipo (Park etal 983090983088983088983097) el grado de madurez del tubeacuterculo (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) y las temperaturasdurante el almacenamiento (Hertog Tijskens amp Hak983089983097983097983095) pueden influir en el crecimiento del brote

En general el contenido de sacarosa y glucosa entodas las variedades estudiadas fue siempre mayor

al de fructosa resultados que concuerdan con losreportados por Rodriacuteguez-Saona y Wrolstad (983089983097983097983095)en la condicioacuten de reposo y de Park et al (983090983088983088983097)en la brotacioacuten de los tubeacuterculos Lo que indica laimportancia de eacutestos azuacutecares sobre todo la sacarosala cual ha sido considerada como la responsable directaen el crecimiento del brote (Debast et al 983090983088983089983089)

Conclusiones

La mayor duracioacuten en el ciclo bioloacutegico de la papa variedad Alpha no repercutioacute en mayor peso total

In addition to sugars (Viola et al 983090983088983088983095) and hormones(Sonnewald amp Sonnewald 983090983088983089983092) the genotype (Parket al 983090983088983088983097) the degree of tuber maturity (Pritchardamp Adam 983089983097983097983090 Sabba et al 983090983088983088983095) and temperaturesduring storage (Hertog Tijskens amp Hak 983089983097983097983095) canaffect sprout growth

In general the content of sucrose and glucose in

all varieties studied was always greater than that offructose results that agree with those reported byRodriacuteguez-Saona and Wrolstad (983089983097983097983095) during dormancyand those of Park et al (983090983088983088983097) during tuber sproutingThis indicates the importance of these sugarsespecially sucrose which has been considered directlyresponsible for sprout growth (Debast et al 983090983088983089983089)

Conclusions

The longer biological cycle of the potato variety Alphadid not result in greater total tuber weight per plantsince it had a similar behavior to that of Gigant and

Vivaldi where the number and weight of commercialtubers per plant were the components that contributedmost to yield

Physiological age (apical dominance multiple sproutingand senescence) did not imply a higher concentrationof soluble sugars in the tubers in this regard Vivaldi

was the variety that showed the highest content ofsucrose and reducing and total sugars

A higher content of reducing and total sugars in thetuber resulted in greater sprout length and dry weight

End of English version

References Referencias

Ap Rees T amp Morrell S (983089983097983097983088) Carbohydrate metabolismin developing potatoes American Journal of Potato

Research 983094983095(983089983090) 983096983091983093-983096983092983095 doi 983089983088983089983088983088983095BF983088983091983088983092983092983090983097983094Cao W amp Tibbitts T W (983089983097983097983092) Phasic temperature

change patterns affect growth and tuberization inpotatoes Journal of the American Society for Horticultural

Science 983089983089983097(983092) 983095983095983093-983095983095983096 Recuperado de httpjournalashspublicationsorgcontent983089983089983097983092983095983095983093fullpdf+htmlCao W amp Tibbitts T W (983089983097983097983093) Leaf emergence on potato

stems in relation to thermal time Agronomy Journal983096983095(983091) 983092983095983092-983092983095983095 doi 983089983088983090983089983091983092agronj983089983097983097983093983088983088983088983090983089983097983094983090983088983088983096983095983088983088983088983091983088983088983089983091x

Davies H V (983089983097983097983088) Carbohydrate metabolism duringsprouting American potato journal 983094983095(983089983090) 983096983089983093-983096983090983095 doi983089983088983089983088983088983095BF983088983091983088983092983092983090983097983091

Debast S Nunes-Nesi A Hajirezaei M R Hofmann J Sonnewald U Fernie A R amp Boumlrnke F (983090983088983089983089) Altering trehalose-983094-phosphate content in transgenic

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1718

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1718

Morales-Fernaacutendez et al

Revista Chapingo Serie Horticultura | Vol XXI nuacutem 2 mayo-agosto 2015

potato tubers affects tuber growth and altersresponsiveness to hormones during sprouting Plant

physiology 983089983093983094(983092) 983089983095983093983092-983089983095983095983089 doi httpdxdoiorg983089983088983089983089983088983092pp983089983089983089983089983095983097983097983088983091

Dogras C Siomos A amp Psomakelis C (983089983097983096983097) Sugarcontent dry matter and sprouting of potato (Solanumtuberosum L) tubers stored at 983094 ordmC and 983089983088 ordmC inrelation to cultivar and area of production Acta

Horticulturae 983090983093983096 983093983092983095-983093983093983092 Recuperado de http wwwactahortorgbooks983090983093983096983090983093983096_983094983090htm

Garciacutea E (983089983097983096983096) Modificacioacuten al Sistema de Clasificacioacuten

Climaacutetica de Koumlppen para Adaptarlo a las Condiciones

de la Repuacuteblica Mexicana Meacutexico UNAM Instituto deGeografiacutea

Haverkort A J Van De Waart M amp Bodlaender K B A(983089983097983097983088) The effect of early drought stress on numbersof tubers and stolons of potato in controlled andfield studies Potato Research 983091983091(983089) 983096983097-983097983094 doi 983089983088983089983088983088983095BF983088983090983091983093983096983089983091983091

Hertog M L A T M Tijskens L M M amp Hak P S (983089983097983097983095)The effect of temperature and senescence on the

acumulation of reducing sugars during storage ofpotato (Solanum tuberosum L) tubers a mathematicalmodel Postharvest Biology and Technology 983089983088(983089) 983094983095-983094983097doi 983089983088983089983088983089983094S983088983097983090983093-983093983090983089983092(983097983095)983096983095983090983095983094-983094

Knowles N R Driskill E P amp Knowles L O (983090983088983088983097)Sweetening responses of potato tubers of differentmaturity to conventional and non-conventional storagetemperature regimes Postharvest Biology and Technology983093983090(983089) 983092983097-983094983089 doi983089983088983089983088983089983094jpostharvbio983090983088983088983096983088983096983088983089983093

Kooman P L Fahem M Tegera P amp Haverkort A J (983089983097983097983094)Effects of climate on different potato genotypes 983090Dry matter allocation and duration of the growthcycle European Journal of Agronomy 983093(983091) 983090983088983095-983090983089983095 doi983089983088983089983088983089983094S983089983089983094983089-983088983091983088983089(983097983094)983088983090983088983091983090-983089

Morales-Fernaacutendez S D Mora-Aguilar R Rodriacuteguez-Peacuterez J E Salinas-Moreno Y Colinas-Leoacuten M amp Lozoya-Saldantildea H (983090983088983089983089) Desarrollo y rendimiento depapa en respuesta a la siembra de semilla-tubeacuterculoinmadura Revista Chapingo Serie horticultura 983089983095(983089) 983094983095-983095983093 doi dxdoiorg983089983088983093983089983093983092rrchsh983090983088983089983089983089983095983088983089983088

Park S W Jeon J H Kim H S Hong S J Aswath Camp Joung H (983090983088983088983097) The effect of size and quality ofpotato microtubers on quality of seed potatoes in thecultivar lsquoSuperiorrsquo Scientia horticulturae 983089983090983088(983089) 983089983090983095-983089983090983097 doi 983089983088983089983088983089983094jscienta983090983088983088983096983088983097983088983088983092

Pritchard M K amp Adam L R (983089983097983097983090) Preconditioning andstorage of chemically immature Russet Burbank andShepody potatoes American Potato Journal 983094983097(983089983090) 983096983088983093-983096983089983093 doi 983089983088983089983088983088983095BF983088983090983096983093983092983089983096983096

Rodriacuteguez-Saona L E amp Wrolstad R E (983089983097983097983095) Influenceof potato composition on chip color quality American

Potato Journal 983095983092(983090) 983096983095-983089983088983094 doi 983089983088983089983088983088983095BF983088983090983096983093983089983093983093983093Sabba R P Bussan A J Michaelis B A Hughes R Drilias

M J amp Glynn M T (983090983088983088983095) Effect of planting and vine-kill timing on sugars specific gravity and skinset in processing potato cultivars American Journal of

de tubeacuterculos por planta ya que eacutesta tuvo uncomportamiento similar a Gigant y Vivaldi en dondeel nuacutemero y peso de tubeacuterculos comerciales por plantafueron los componentes que maacutes contribuyeron alrendimiento

La edad fisioloacutegica (dominancia apical brotacioacutenmuacuteltiple y senescencia) no implicoacute mayor concentracioacuten

de azuacutecares solubles en el tubeacuterculo y en eacutestas Vivaldifue la variedad que registroacute el mayor contenido desacarosa azuacutecares reductores y totales

El contenido de azuacutecares reductores y totales en eltubeacuterculo repercutieron en mayor longitud y peso secodel brote

Fin de la versioacuten en espantildeol

Potato Research 983096983092(983091) 983090983088983093-983090983089983093 doi 983089983088983089983088983088983095BF983088983090983097983096983094983090983095983088

Sinha N K Cash J N amp Chase R W (983089983097983097983090) Differencesin sugars chip color specific gravity and yield ofselected potato cultivars grown in Michigan American

potato journal 983094983097(983094) 983091983096983093-983091983096983097 doi 983089983088983089983088983088983095BF983088983090983096983095983095983092983096983097Sonnewald S amp Sonnewald U (983090983088983089983092) Regulation of potato

tuber sprouting Planta ( 983090983091983097) 983090983095-983091983096 doi 983089983088983089983088983088983095s983088983088983092983090983093-983088983089983091-983089983097983094983096-z

Sowokinos J R (983090983088983088983089) Biochemical and molecular controlof cold-induced sweetening in potatoes American

Journal of Potato Research 983095983096(983091) 983090983090983089-983090983091983094 doi 983089983088983089983088983088983095BF983088983090983096983096983091983093983092983096

Stark J C amp Love S l (983090983088983088983091) Tuber quality In Stark J C ampLove S L (Eds) Potato production systems (pp 983091983090983097-983091983092983091) Idaho Center of Potato Research and Education

Statistical Analysis System (SAS Institute) (983090983088983088983092) SASSTAT983097983089 userrsquos guide Cary NC USA Autor

Struik P C Haverkort A J Vreugdengil D Bus C Bamp Dankert R (983089983097983097983088) Manipulation of tuber-sizedistribution of a potato crop Potato Research 983091983091(983092)983092983089983095-983092983091983090 doi 983089983088983089983088983088983095BF983088983090983091983093983096983088983089983097

Suttle J C (983090983088983088983092) Physiological regulation of potato tuberdormancy American Journal of Potato Research 983096983089(983092)983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983095

Thompson A L Love S L Sowokinos J R Thornton MK amp Shock C C (983090983088983088983096) Review of the sugar end

disorder in potato (Solanum tuberosum L) American Journal of Potato Research 983096983093(983093) 983091983095983093-983091983096983094 doi 983089983088983089983088983088983095s983089983090983090983091983088-983088983088983096-983097983088983091983092-983090

Van Es A amp Hartmans Y (983089983097983096983089) Sugars and starch duringtuberization storage and sprouting In Rastovski

A amp Van Es A (eds) Storage of potatoes (pp 983096983090-983097983096) Wageningen The Netherlands Pudoc

Viola R Pelloux J van der Ploeg A Gillespie T MarquisN Roberts A G amp Hancock R D (983090983088983088983095) Symplasticconnection is required for bud outgrowth followingdormancy in potato (Solanum tuberosum L) tubers

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092

7232019 Crecimiento rendimiento y contenido de azuacutecares a diferente edad fisioloacutegica del tubeacuterculo de papa

httpslidepdfcomreaderfullcrecimiento-rendimiento-y-contenido-de-azucares-a-diferente-edad-fisiologica 1818

983089983092983094 Growth yield and sugar

Plant Cell Environment (983091983088) 983097983095983091ndash983097983096983091 doi 983089983088983089983089983089983089j983089983091983094983093-983091983088983092983088983090983088983088983095983088983089983094983097983090x

Vreugdenhil D (983090983088983088983092) Comparing potato tuberization andsprouting Opposite phenomena American journal of

potato research 983096983089(983092) 983090983095983093-983090983096983088 doi 983089983088983089983088983088983095BF983088983090983096983095983089983095983094983097 Walworth J L amp Carling D E (983090983088983088983090) Tuber initiation and

development in irrigated and non-irrigated potatoes American journal of potato research 983095983097(983094) 983091983096983095-983091983097983093 doi

983089983088983089983088983088983095BF983088983090983096983095983089983094983096983091

Wiersema S G (983089983097983096983093) Desarrollo Fisioloacutegico de Tubeacuterculos-

Semilla de Papa Centro Internacional de la Papa LimaPeruacute

Wolf S (983089983097983097983091) Effect of leaf age on photosynthesis carbontransport and carbon allocation in potato plants Potato research 983091983094(983091) 983090983093983091-983090983094983090 doi 983089983088983089983088983088983095BF983088983090983091983094983088983093983091983092