Tesis Doctoral

Acumulación y distribución de
macronutrientes minerales en dos

cultivares de Solanum tuberosum L.
en diferentes ambientes del altiplano
Cundi-boyacense
Manuel Iván Gómez Sánchez
Universidad Nacional de Colombia

Facultad de Ciencias Agrarias
Bogotá, Colombia
2018
Acumulación y distribución de
macronutrientes minerales en dos
cultivares de Solanum tuberosum L.
en diferentes ambientes del altiplano
Cundi-boyacense
Manuel Iván Gómez Sánchez
Tesis de doctorado presentada como requisito parcial para optar al título de:
Doctor en Ciencias Agrarias
Director:
Ph.D. Stanislav Magnitskiy
CODIRECTOR:
Ph.D Luis Ernesto Rodríguez
Línea de Investigación:
Fisiologia Vegetal
Universidad Nacional de Colombia

Facultad de Ciencias Agrarias
Escuela de Posgrados
Bogotá, Colombia
2018
Dedicatoria
A la persona más importante que me dio la
vida, la salvación, la sabiduria, la sanidad, la
ciencia y el conocimiento a mi Señor y Dios
y Salvador Jesucristo gracias por ser todo
en mi vida.
A mi esposa, ayuda idónea, hermosa sabia y

amada Mariana Salamanca; a mi
primogénito caballero Emanuel David, a mi
hija que soñe Laura Valentina; a mi
hermosa, tierna y amada hija Marianne
Andressa y al hijo de mi vejez mi pequeño
caballero Ivan Yeshua. Gracias por su
paciencia y fé que junto a sus oraciones
permitieron que llegara a esta meta, los amo.
A mis padres, hermanos y mi padre espiritual

Eduardo Dosantos que me han dado ejemplo
de integridad, paciencia, perseverancia,
sacrificio y amor. Gracias por creer en mi.
“Como el Padre me ha amado, así también

yo os he amado; permaneced en mi amor”
Jesus de Nazaret tu Salvador Juan 15:9

“El Presidente de Tesis, el Consejo de
Jueces de Tesis y el Consejo
examinador no serán responsables de
las ideas emitidas por el candidato”.
(artículo 152 de los Estatutos de la
Universidad Nacional de Colombia, Acuerdo
66 de 1939).
Agradecimientos
Agradezco a mis directores: Dr Stanislav Magnitskiy (Unal-Bogotá) y Dr Luis Ernesto
Rodriguez (Unal-Bogotá) por su dirección idónea, seguimiento y aportes científicos en la
planeación, realización y culminación de los estudios doctorales.
Agradezco a la Facultad de Ciencias Agrarias de la Unal-Bogotá y el soporte dado en

los laboratorios de fisiología vegetal, equipos e instalaciones y en especial a los
profesores e investigadores: Dra Liz Moreno, Dr Herman Restrepo, Dr Victor Flores y Dr
Carlos Ñústez (Unal-Bogotá); Dr Walter Osorio (Unal-Medellin); Dr Raul Jaramillo (IPNI-
Ecuador); Dr Enrique Combatt (Ucordoba-Monteria) y Dr Pedro Almanza (UPTC-Tunja)
por sus apoyos científicos en la revisión de los seminarios de investigación y aportes
académicos para la construcción de la investigación de la tesis doctoral.
Agradezco al Dr. Enrique Darghan y al I.A. Johan Urquijo de la Unal-Bogotá por los
valiosos aportes en el análisis estadístico y el desarrollo de modelos estadísticos
novedosos para la interacción de factores genotipo x ambiente.
Agradezco a la empresa privada Ingeplant Ingenieria en Nutrición de Cultivos SAS

por el epoyo en tiempo para los estudios doctorales y la financiación económica del 80%
de la investigación doctoral, además, resalto el soporte loable de campo y laboratorio que
me brindo el grupo técnico y de investigación de esta empresa con los ingenieros
agrónomos: Andrea Barragan, Liliana Arevalo, Elias Silva y Paola Torres en cada una de
la fases de la investigación. También una mención especial a Fedepapa por la
financiación del 20% de la investigación en la primera etapa.
Agradezco y exalto especialmente el soporte técnico y apoyo logístico en cultivo del

Ingeniero Walter Guzmán y su empresa Biogenetica-Chocontá y a los agricultores
Ricardo Rojas y Giovanny Pulido en Facatativá y Carlos Acero en Subachoque.
Resumen y Abstract IX
Resumen
Los estudios de nutrición mineral en cultivares de papa del Grupo Andigena en Colombia, se
han enfocado principalmente a respuestas de la fertilización N, P y K. Sin embargo, no se ha
cuantificado el comportamiento de estos nutrientes en la planta con relación a las diferentes
etapas fenológicas y su distribución en los diferentes órganos, ni su relación con
componentes fisiológicos de crecimiento para su modelamiento. Por ello en la presente
investigación, bajo ambientes contrastantes de evaluación (suelos de baja y alta fertilidad),
dos ciclos de producción, dos cultivares (Diacol Capiro [Capiro] y Pastusa Suprema
[Suprema]) y fertilización variable, se buscó: (i) caracterizar la distribución de macronutrientes
en órganos de la planta; (ii) determinar la acumulación de N, P y K en diferentes etapas del
cultivo (iii) y establecer a partir de relaciones alométricas la acumulación de nutrientes
minerales, variables fisiológicas de crecimiento y uso eficiente de los nutrientes. Se
determinó que la curva de dilución crítica para cada macronutriente en Capiro es Nc = 6,23
-0,319 -0,198 -0,269 -0,327
W , Pc = 0,523 W y Kc = 9,02 W , y para Suprema de Nc= 6,74 W , Pc=
-0,186 -0,1353
0,536 W y Kc= 6,585 W , observandose mejor robustez con peso seco total (W)
con respecto al modelo obtenido con índice de área foliar. Se observó además significancia
estadística en la interacción genotipo x localidad con mejor ajuste en el modelo de consumo
nutricional para suelos de baja fertilidad para Suprema, con Nr = 68,13 W0,504, Pr = 6,72
W0,779 y Kr = 63,93 W0,776, donde se expresa el mayor potencial productivo y altos índices de
cosecha por nutriente con ICN = 0,55-0,69, ICP = 0,75-0,8 y ICK = 0,62, mientras que Capiro
muestra una mayor adaptación en ambos tipos de suelos, con una mejor conversión de
asimilados por su uso eficiente de nutrientes en el tubérculo (UENt) y con Nr = 56,38 W0,58 y
Pr = 4,26 W0,786 en baja fertilidad, y Kr = 79,52 W0,79 en alta fertilidad. Se estableció el índice
de nutrición (INN) con mejor ajuste en Capiro entre 0,25-1,32, en contraste, para Suprema se
evidenció una respuesta entre nula a marginal a la fertilización y consumo de lujo de N (INN
1-1,5) que fue corroborado con inicadores negativos en el UENt, una menor eficiencia de
traslocación (tanto en N, P y K) y una mayor acumulación de NO 3ˉ y K+ en savia bajo suelos
de alta fertilidad. Lo anterior permitirá realizar diagnósticos nutricionales oportunos y así
pronosticar un manejo más específico de estos nutrientes por cultivar y diferentes tipos de
suelo para alcanzar altos rendimientos y sostenibilidad en el cultivo de papa en la región
Andina de Colombia.
Palabras clave: uso eficiente de nutrientes, relaciones alométricas, consumo de lujo,

eficiencia de traslocación, papa del grupo Andigena.
X Resumen y Abstract
Abstract
Studies of mineral nutrition in potato crop of Group Andigena in Colombia have been
focused mainly on plant responses to fertilization with N, P, and K. However, the behavior
of macronutrients in these plants has not been quantified with respect to the phenological
stages and macronutrient allocation to the different organs or their relationship with the
physiological components of growth. For this reason, the research was undertaken in
contrasting environments and two production cycles using two cultivars (Diacol Capiro
and Pastusa Suprema) under variable fertilization conditions pursuing the following
objectives to (i) characterize the distribution of N, P, and K in organs of the plant; (ii)
determine the accumulation of macronutrients in different stages of crop growth, and (iii)
establish from allometric relationships the accumulation of mineral nutrients, physiological
variables of growth, and efficient use of nutrients. Critical dilution curves based on total
-0.319 -0.198
dry biomass (W) were determined for Capiro: Nc = 6.23 W ; Pc = 0.523 W ; Kc =
9.02 W -0.269, and for Suprema: Nc = 6.74 W -0.327
; Pc = 0.536 W -0.186; Kc = 6,585W -0.1353
,
observing better robustness with respect to the models obtained with leaf area index.
Genotype x locality interaction was identified with the best fit of the uptake model in low
fertility soils for Suprema (Nr = 68.13 W0,504, Pr = 6.72 W0.799 and Kr = 63.93 W0,776),
where it expressed the highest productive potential and high harvest index HIN (0.55-
0.69), HIP (0.75-0.8) and HIK (0.62), while Capiro showed a better adaptation to both
types of soils with a better efficient use of nutrients in the tuber in UENt and with Nr =
56.38 W0.58 and for Pr = 4.26 W0.786 (low fertility) and for Kr = 79.52 W 0.79 (high fertility).
The nutrinte index-INN (0.25-1.32) with better adjustment in Capiro was established, in
contrast for Suprema it is evidenced the null to marginal response to the fertilization and
luxury uptake of N (INN 1-1,5) that was corroborated with the negative indices in UENt, a
lower efficiency of translocation (EtN, EtP, and EtP) and a greater accumulation of NO 3ˉ
and K+ in sap under high fertility soils. The above data facilitate nutritional diagnostics
and, thus, prediction a more specific management of these nutrients by the crop and soil
type to achieve high yields and sustainability in potato production in the Andean region of
Colombia.
Key words: efficient use of nutrients, allometric relationships, luxury consumption,

translocation efficiency, potato of Group Andigena.
Contenido XI
Contenido
Pág.
Lista de figuras .............................................................................................................. XIV

Lista de tablas .............................................................................................................. XVII
Lista de Símbolos y abreviaturas ................................................................................... XIX
Introducción.......................................................................................................................1
1. Critical Dilution Curve for N, P, and K and Leaf Area Index in Potato (Solanum
tuberosum L., Group Andigena) ...................................................................................... 15
1.1 Abstract .............................................................................................................15
1.2 Resumen ...........................................................................................................16
1.3 Introduction .......................................................................................................17
1.4 Materials and methods ......................................................................................19
1.4.1 Field experiments .......................................................................................... 19
1.4.2 Analytical methods......................................................................................... 22
1.4.3 Data analysis ................................................................................................. 22
1.5 Results and discussion ................................................................................. ….24
1.5.1 Critical Dilution Curves for Nc, Pc, and Kc ..................................................... 24
1.5.2 Nutrition index and optimal fertilization dose .................................................. 30
1.6 Conclusions.......................................................................................................34
1.7 Bibliografía ........................................................................................................34
2. Uptake and partition of N, P and K in potato (Solanum tuberosum L. Group.
Andigena) ……………………………………………………………………………………..39
2.1 Abstract .............................................................................................................39
2.2 Resumen ...........................................................................................................40
2.3 Introducción.......................................................................................................41
2.4 Materials and methods ......................................................................................42
2.4.1 Experimental locations and conditions ........................................................... 42
2.5 Analytical methods ............................................................................................45
XII Acumulación y distribución de macronutrientes minerales en dos cultivares de
Solanum tuberosum L. en diferentes ambientes del altiplano Cundi-boyacense
2.5.1 Statistical analysis ..........................................................................................45

2.6 Results.............................................................................................................. 46
2.6.1 Total uptake of N, P, and K and nutrient harvest indexes ...............................46
2.6.2 Efficiency of translocation and extraction of N, P, and K by tubers .................52
2.7 Discussion ........................................................................................................ 55
2.7.1 Total uptake of N, P, and K and nutrient harvest indexes ...............................55
2.7.2 Efficiency of translocation and extraction of N, P, and K by tubers .................58
2.8 Conclusions ...................................................................................................... 59
2.9 Bibliografía ........................................................................................................ 60
3. Accumulation of N, P, and K in the tubers of potato (Solanum tuberosum L. spp.
andigena) under contrasting soils of the Andean region of Colombia ...............................65
3.1 Abastract .......................................................................................................... 65
3.2 Resumen .......................................................................................................... 66
3.3 Introduction ....................................................................................................... 67
3.4 Materials and methods ...................................................................................... 68
3.5 Results and discussion ..................................................................................... 71
3.5.1 Accumulation of N, P and K in tuber phenology ..............................................71
3.5.2 Accumulation of N, P, and K among cultivars .................................................76
3.6 Conclusions ...................................................................................................... 79
3.7 Literature cited .................................................................................................. 80
4. Potential yield and efficiency of N and K uptake in tubers of cvs. Diacol Capiro and
Pastusa Suprema (Solanum tuberosum subsp. andigena)...............................................85
4.1 Abstract ............................................................................................................ 85
4.2 Resumen .......................................................................................................... 86
4.3 Introduction ....................................................................................................... 87
4.4 Materials and methods ...................................................................................... 88
4.4.1 Location and soils...........................................................................................88
4.4.2 Plant sampling and analysis ...........................................................................90
4.4.3 Efficiency use and recovery of mineral nutrients by tubers .............................91
4.4.4 Statistical analysis ..........................................................................................91
4.5 Results and discussion ..................................................................................... 92
4.5.1 Yield, harvest index, and dry weight of tubers.................................................92
4.5.2 Efficiency in the nutrient use by tubers ...........................................................97
4.6 Conclusions .................................................................................................... 102
4.7 Literature cited ................................................................................................ 103
Introducción XIII
5. Diagnóstico de K+ y NO3ˉ en savia para determinar el estado nutricional en papa

(Solanum tuberosum L. subsp. andigena) ..................................................................... 107
5.1 Resumen .........................................................................................................107
5.2 Abstract ...........................................................................................................108
5.3 Introducción.....................................................................................................109
5.4 Materiales y métodos ......................................................................................111
5.5 Resultados y discusión ....................................................................................114
5.5.1 K+ y N-NO3ˉ en savia por etapa fenológica y cultivar .................................... 114
5.5.2 N-NO3ˉ y K+ savia y su relación con peso seco y rendimiento ...................... 120
5.6 Conclusiones...................................................................................................123
5.7 Referencias bibliográficas ...............................................................................124
6. Conclusiones.......................................................................................................... 129
7. Recomendaciones.................................................................................................. 131
8. Anexo A: Anovas de rendimiento, eficiencia de traslocación, índices de cosecha y
covariables ……………………………………………………………………………………133
9. Anexo B: Coeficientes de consumo, coeficientes de traslocación e índice de cosecha
de nutriente ……………………………………………………………………………………139
10. Anexo C: Componentes principales y análisis de diseño en medidas repetidas de
acumulación de biomasa, nutrientes y uso eficiente de nutrientes en el tubérculo ........ 141
Contenido XIV
Lista de figuras
Pág.
Figure 1-1. Critical dilution curves for N (A), P (B), and K (C) in cvs. Capiro ( ) and
Suprema ( ) in two production cycles (2013-2016) under non-limiting conditions of
fertilization in soils of contrasting fertility. Means consisted of n = 13 data per phenological
stage and cultivar. Coefficients a and b of potential allometric function were based on total
dry biomass W (equation 1). ........................................................................................... 25
Figure 1-2. Critical dilution curves for N (A), P (B) and K (C) based on LAI in cvs. Capiro
( ) and Suprema ( ) in two production cycles (2013-2016) under non-limiting conditions
of fertilization in soils of contrasting fertility. n = 65 data per cultivar. Coefficients a and b
of allometric function were based on LAI (equation 2). .................................................... 26
Figure 1-3. Relationship between relative yield (RY) and nitrogen nutrition index (NNI) (A)
in cvs. Capiro and Suprema at tuber filling (125-150 dap) and differential optimum
fertilizer dose (dODF) (150-160 dap) (B) for cvs. Capiro ( ) and Suprema ( ) in two
production cycles in soils contrasting in fertility. ***P <0.001; *P < 0.05; ns, not significant.
....................................................................................................................................... 31
Figure 2-1. Average total uptake of N, P, and K in cvs. Capiro (left) and Suprema (right)
as a function of total dry biomass (W) in soils of high (Andic Eutrudepts, Facatativa) and
low (Humic Dystrudepts, Chocontá) fertility and two production cycles under non-limiting
conditions of fertilization. **Indicates significant values of coefficients b with respect to soil
fertility at a probability level of 0.05. ns, non-significant model. ....................................... 47
Figure 2-2. Potential yield (PYt) in cvs. Capiro and Suprema cultivated in soils of low
(Humic Dystrudepts, Chocontá) or high (Andic Eutrudepts, Facatativá) fertility at 150-160
dap under non-limiting conditions of fertilization in two production cycles. P<0.05 for
fertilization x location x cultivar. Error bars indicate standard errors. ............................... 48
Figure 2-3. Harvest index (HI) in cvs. Capiro and Suprema cultivated in soils of low
(Humic Dystrudepts, Chocontá) or high (Andic Eutrudepts, Facatativá) fertility under non-
limiting conditions of fertilization in two production cycles. P<0.05 for location x cultivar x
cycle. Means followed by the different letters are significantly different (LSD, P<0.05).
Error bars indicate standard errors. ................................................................................. 48
Figure 2-4. Harvest index for N (NHI) (A), P (PHI) (B), and K (KHI) (C) in cvs. Capiro and
Suprema cultivated on Humic Dystrudepts (Chocontá) and Andic Eutrudepts (Facatativá).
P<0.001 for location x cultivar x cycle under non-limiting conditions of fertilization. Means
1. Critical Dilution Curve for N, P, and K and Leaf Area Index in Potato XV
(Solanum tuberosum L., Group Andigena)
followed by the different letters are significantly different (LSD, P<0.05). Error bars
indicate standard errors. ..................................................................................................50
Figure 2-5. Average translocation efficiency of N (EtN), P (EtP), and K (EtK) in cv. Capiro
(circle) and cv. Suprema (square) in two production cycles under non-limiting conditions
of fertilization in contrasting soils. ** Significant differences of coefficients b are shown
with respect to the interaction cultivar x location at a probability level of P <0.05. ............53
Figure 3-1. Accumulation of N P K in tubers during the growing season in andigena
potato, cvs. Suprema (left) and Capiro (right) in Typic Hapludands (Subachoque, ♦);
Humic Dystrudepts (Chocontá, ■) and Andic Eutrudepts (Facatativá, ▲) with fertilization
(fer1) in the altiplano Cundiboyacense. Dap, day after planting. Error bars indicate
standard error. .................................................................................................................73
Figure 3-2. Relationship between the uptake in tubers of N (A), P (B) and K (C) (kg ha -1)
yield (FWT), and cvs. Capiro and Suprema in soils of Cundiboyacense plateau. ** The
value of coefficients b were significant at the 0.05 probability level. ns, not significant the
value coefficients b. .........................................................................................................77
Figure 4-1. Yield (FWt) of cvs. Capiro and Suprema at the phenological stage V
(maximum tuber filling and maturation) in the absence of fertilization F0, with respect to
the balanced fertilization by location: F1s, Typic Hapludands (Subachoque); F1ch, Humic
Dystrudepts (Chocontá), and F1f, Andic Eutrudepts (Facatativá). P<0.001 for fertilization
(location * cultivar) ...........................................................................................................92
Figure 4-2. Harvest index (HI) in cvs. Diacol Capiro and Pastusa Suprema in the absence
of fertilization Fer0, with respect to balanced fertilization by location: F1s, Typic
Hapludands (Subachoque); F1ch, Humic Dystrudepts (Chocontá), and F1f, Andic
Eutrudepts (Facatativá). P<0.001 for location*cultivar. ....................................................94
Figure 4-3. Tuber dry weight (DWt) in cvs. Diacol Capiro and Pastusa Suprema at four
phenological stages (II, start of tuberization; III, maximum tuberization-start of filling; IV,
filling of tuber; V, maximum filling and maturation) in the absence of fertilization F0, with
respect to balanced fertilization by location for soils Typic Hapludands (Subachoque),
F1s; Humic Dystrudepts (Chocontá), F1ch and Andic Eutrudepts (Facatativá), Fer1f;
P<0.001 for fertilization (location * cultivar). .....................................................................96
Figure 4-4. Efficiency of recovery of N and K in tubers (RFt), kg nutrient extracted per 100
kg nutrient applied in balanced fertilization, cvs. Diacol Capiro and Pastusa Suprema on
contrasting soils of the Andean region-Colombia. P<0.001 in N and K for location *
cultivar. ............................................................................................................................97
Figure 4-5. EPt of N and K (kg of tuber harvested per kg of nutrient extracted) in ‘Diacol
Capiro’ and ‘Pastusa Suprema’ in contrasting soils of the Andean region-Colombia.
P<0.001 in N and K for location * cultivar. .......................................................................98
Figure 4-6. Efficient use of N and K in tubers of balanced fertilization, NUEt (kg of dry
matter of the tuber per kg of nutrient extracted) in cvs. Diacol Capiro and Pastusa
XVI Acumulación y distribución de macronutrientes minerales en dos cultivares de
Suprema in Typic Hapludands, Subachoque, Andic Eutrudepts, Facatativá, and Humic

Dystrudepts, Chocontá in Andean region-Colombia. ..................................................... 100
Figura 5-1. Variación de K+ en savia fresca de tallo medida en campo en el ciclo del
cultivo de ‘Diacol Capiro’ (A) y ‘Pastusa Suprema’ (B) en respuesta a la fertilización en
suelos de alta fertilidad de la Sabana de Bogotá. Letras diferentes entre tratamientos
para cada etapa fenológica presenta diferencias estadísticas significativas (Tukey,
P<0,05). ........................................................................................................................ 115
Figura 5-2. Variación de NO3ˉ en savia fresca de tallo medida en campo en el ciclo de
para cada época presenta diferencias estadísticas significativas, Tukey, P<0,05. ........ 116
Figura 5-3. Relación entre área foliar y el contenido de NO 3ˉ (A) y K+ (B) en savia fresca
de tallo medida en campo hasta etapa III (90-100 dds), floración, máxima tuberización e
inicio de llenado en papa ‘Diacol Capiro’ en respuesta a la fertilización en suelos de alta
fertilidad de la Sabana de Bogotá. ** Modelo altamente significativo (P<0,01); * Modelo
significativo (P<0,05). .................................................................................................... 118
Figura 5-4. Relación entre las concentraciones K + (A) y N-NO3ˉ (B) en savia de tallo
medida en campo y el rendimiento, Pft medido desde etapa de floración a maduración
para ‘Diacol Capiro’ (cuadro) y ‘Pastusa Suprema’ (rombo) en suelos de alta fertilidad de
la Sabana de Bogotá..................................................................................................... 121
Figura 5-5. Respuesta en rendimiento en ‘Diacol Capiro’ (A) y ‘Pastusa Suprema’ (B) a la
fertilización y su relación con concentración de K + y N-NO3ˉ en savia de tallo en suelos de
alta fertilidad de la Sabana de Bogotá. .......................................................................... 122
Figura 10-1. Componentes principales de variable y acumulación de nutrientes en
tubérculos. .................................................................................................................... 141
Contenido XVII
Lista de tablas
Pág.
Table 1-1. Environmental and soil fertility characteristics in studied locations. .................20
Table 1-2. Doses of mineral nutrients in fertilizer treatments. ..........................................21
Table 1-3. Allometric relationships and critical dilution coefficients for N, P, and K based
on total dry biomass W (above-ground organs and tubers) for cvs. Capiro and Suprema
as compared with cultivars of Group Chilotanum. The data for cvs. Capiro and Suprema
were obtained under non-limiting conditions of fertilization in soils of contrasting fertility. 27
Table 1-4. Allometric relations and critical dilution coefficients for N, P, and K based on
leaf area index (LAI) for cvs. Capiro and Suprema under non-limiting conditions of
fertilization in soils of contrasting fertility. .........................................................................29
Table 1-5. Quadratic model for yield and balanced dose of fertilizer; c: quadratic
coefficient; b: linear coefficient; a: intercept, R2: coefficient of determination. MY –
maximum yield; ODF – optimal dose of fertilizer to achieve maximum yields. .................32
Table 1-6. Average yield of tubers (Mg ha –1) harvested at 150-160 dap for cvs. Capiro and
Suprema and statistical significance of mean square of factors and their interactions. ....33
Table 2-1. Characteristics of climate and soils in experimental locations. ........................43
Table 2-2. Contribution of mineral nutrients through fertilizer applications. ......................44
Table 2-3. Prognosis of total uptake and extraction by tubers for N, P and K in cvs. Capiro
and Suprema at different yields in soils of high yield potential of the Colombian Andean
region. .............................................................................................................................52
Table 2-4. Extraction of N, P, and K by tubers per growth stage for cvs. Suprema and
Capiro. The values are estimates from translocation efficiency of N (EtN), P (EtP), and K
(EtK) with a projected yield of 70 Mg ha−1. .......................................................................54
Table 3-1. Edaphoclimatic conditions in the locations of the study...................................69
Table 3-2. Doses of mineral nutrients applied with fertilization in the studied locations. ...70
Table 4-1. Environmental and soil fertility characteristics at the study sites. ....................89
Table 4-2. Doses of mineral nutrients applied with fertilizers in the study sites. ...............90
Tabla 5-1. Niveles de NO3ˉ y K+ en savia de peciolo de papa en diferentes estados
fenológicos evaluados con medidores de ion selectivo. .................................................110
XVIII Acumulación y distribución de macronutrientes minerales en dos cultivares de
Tabla 5-2. Propiedades químicas del suelo en el sitio de evaluación. ........................... 112
Tabla 5-3. Aporte de nutrientes minerales en los tratamientos con fertilización edáfica
(dosis en kg ha-1). ......................................................................................................... 113
Tabla 8-1. Anovas e interacciones de factores para consumo y eficiencia de traslocación
de N, P y K en cuatro etapas del cultivo e índice de cosecha de nutrientes 150 dds..... 133
Tabla 8-2. Análisis de regresión de covariables de suelo y clima respecto a las variables
evaluadas. .................................................................................................................... 135
Tabla 8-3. Anovas e interacciones de factores en las variables rendimiento. ................ 137
Tabla 8-4. Anovas e interacciones de factores en las variables rendimiento por cultivar.
..................................................................................................................................... 137
Tabla 8-5. Anovas e interacciones de factores para índice de cosecha por cultivar. ..... 138
Tabla 9-1. Coeficientes de consumo crítico de N, P y K a partir de la biomasa total para
Capiro y Suprema. ........................................................................................................ 139
Tabla 9-2. Coeficientes a y b de eficiencia de traslocación de N, P y K para Capiro y
Suprema sin fertilización (0) y bajo condiciones optimas de fertilización (1). ................. 140
Tabla 10-1. Algoritmo de componentes principales de variables fisiológicas de
crecimiento, consumo y concentración de nutrientes para análisis exploratorio y reducir la
dimensionalidad del conjunto de variables. ................................................................... 142
Tabla 10-2. Algoritmo para las variables fisiológicas de crecimiento (biomasa seca, área
foliar, consumo de nutrientes, uso eficiente de nutrientes) diseño en medidas repetidas
factorial incompleto en parcelas subdivididas-modelo mixto. Los factores entre sujetos
son anidados, pues la naturaleza de la matriz es incompleta. ....................................... 146
Tabla 10-3. Análisis estadístico MANOVA diseño en medidas repetidas (factorial
incompleto en parcelas subdivididas-modelo mixto inter sujetos para los componentes
que reunía la variables de mayor significancia . ............................................................ 148
incompleto en parcelas subdivididas-modelo mixto intrasujetos para los componentes que
reunía la variables de mayor significancia. .................................................................... 149
Tabla 10-5. Matriz de correlación de Pearson de los componentes de alta significancia.
..................................................................................................................................... 149
Tabla 10-6. Coeficiente b del modelo lineal de la extracción de nutrientes N, P y K (
kg/ha) respecto al rendimiento. ..................................................................................... 150
Contenido XIX
Lista de Símbolos y abreviaturas

Abreviatura Término
AF Área foliar
a Concentración del nutriente cuando el total de la biomasa es ≤ 1 t ha -1
b Coeficiente de dilución
Capiro Cultivar Diacol Capiro
CDC Curvas de dilución crítica
cmol Centimol
cv. (s) Cultivar (es)
DAF Duración del área foliar
dds Días después de siembra
dOFR Diferencia de la dosis óptima de fertilización
DWT Dry weight of tubers/Peso seco de tubérculos
EE Error estándar
EFN Eficiencia fisiológica del nutriente
Accumulated nutrients per tuber yield/Eficiencia de nutrientes para la

EPt
producción de tubérculos
EtK Eficiencia de traslocación de potasio en el tubérculo
EtN Eficiencia de traslocación de nitrógeno en el tubérculo
EtP Eficiencia de traslocación de fósforo en el tubérculo
EUN Eficiencia de utilización del nutriente

XX Acumulación y distribución de macronutrientes minerales en dos cultivares de
FWT Fresh weight of tubers/Peso fresco de tubérculos
ha hectárea
HEI harvest extraction index
IAF Índice del área foliar
IC/HI Índice de cosecha/Harvest index
IEC/EI Índice de extranción/Rate of nutrient extraction
INN Índice de nutrición
ISE Método de electrodo selectivo de iones
Kc Curva de dilución de potasio
Kr Consumo de potasio
MO Materia orgánica
Nc Curva de dilución de nitrógeno
Nr Consumo de nitrógeno
Nutrient use efficiency in tubers/ Eficiencia de uso de nutrientes en

NUEt-UENt
tubérculos
PFt Rendimeinto potencial
Pc Curva de dilución de fósforo
Pr Consumo de fósforo
PS Peso seco
OFR Dosis óptima de fertilización
Recuperación de nutrientes del fertilizante por el tubérculo/Rates of

RFt
nutrient recovery efficiency
RR Rendimientos relativos
Subsp., spp Subespecie
Suprema Cultivar Pastusa Suprema
t tonelada
1. Critical Dilution Curve for N, P, and K and Leaf Area Index in Potato XXI
W Biomasa seca total

Introducción
La papa (Solanum tuberosum L.) ha sido consumida por la humanidad desde hace miles
de años y presenta alta versatilidad y adaptabilidad a diferentes condiciones ambientales
(Birch et al., 2012; De Jong, 2016; Hardigan et al., 2017). Actualmente se cultiva en 149
países en latitudes que van desde los 65°N hasta los 50°S y altitudes que varían desde
el nivel del mar hasta los 4.000 m, con una producción anual aproximada de 330 millones
de toneladas (Gebhardt, 2013). Ocupa un lugar importante en la agricultura, economía,
seguridad alimentaria e industria considerándose un cultivo primario básico y
actualmente el tercer cultivo de mayor consumo en el mundo después del maíz y el trigo
(Birch et al., 2012; De Jong, 2016; Li et al., 2013).
Se espera que la población mundial llegue a 9,4 billones de personas en el año 2050
(FAO, 2013) y en estas condiciones será necesario incrementar los rendimientos de los
cultivos y la eficiencia de la producción para lograr un incremento constante, para suplir
la demanda de alimentos. Se estima que los sistemas de producción de maíz, arroz y
papa están funcionando en promedio a 40 y 65% del potencial de rendimiento y que se
necesitan de un incremento entre el 70-80% del potencial de rendimiento para lograr
satisfacer las demandas en los próximos 30 años (Stewart, 2007; Dooberman, 2007;
IPNI, 2009; FAO, 2013).
Para lograr lo anterior, en el ámbito nacional es necesario establecer estrategias que

permitan obtener altos rendimientos, que integren también la conciencia ambiental y la
rentabilidad del agricultor (Bruulsema et al., 2008). El manejo óptimo de nutrientes y un
incremento en la eficiencia de uso serán componentes importantes para lograr este
objetivo (Stewart, 2007; IPNI, 2009).
Para el año 2016, en Colombia se logró una producción estimada de 2,623,700 t ha -1 en

126.100 ha, y los departamentos de Cundinamarca y Boyacá generaron el 76% de la
producción de papa (Riascos, 2016). Diacol Capiro (en adelante Capiro) y Pastusa
Suprema (Suprema) son considerados los cultivares de mayor importancia económica y
representan el 80% del área cultivada en el país para consumo fresco y procesamiento
industrial (Ñústez, 2011).
2 Acumulación y distribución de macronutrientes minerales en dos cultivares de
Desde el punto de vista de la productividad agrícola, la nutrición mineral es uno de los

factores más limitantes en el crecimiento y por lo tanto, es necesario el manejo óptimo de
nutrientes y fertilización al sistema productivo de papa (Wensterman, 2005; Villamil,
2005; Schilling et al., 2016). El conocimiento de la nutrición mineral vegetal permite a la
planta proporcionar el nutriente correcto, la cantidad necesaria, el lugar indicado y el
momento oportuno para conseguir los óptimos fisiológicos en la producción y calidad de
los cultivos en un entorno sostenible y rentable (Wensterman, 2005; IPNI, 2009).
La fisiología de la nutrición mineral permite entender el funcionamiento de la planta y

según Villamil (2005) y Marshner (2012), el estudio de la nutrición mineral comprende: (i)
clasificación de los nutrientes minerales; (ii) mecanismos de absorción de iones por las
células de la raíz; (iii) transporte a corta distancia; (iv) transporte a larga distancia en el
xilema y floema y sus regulaciones; (v) toma y distribución de elementos minerales por
las hojas y órganos; (vi) crecimiento y su relación con la nutrición; (vii) rendimiento y
relación fuente-demanda; (viii) rendimiento y nutrición mineral; (ix) funciones de
macronutrientes y micronutrientes; (x) deficiencias y toxicidad de macronutrientes y
micronutrientes y (xi) relación entre nutrición mineral y enfermedades.
El crecimiento vegetal, tanto en los sistemas naturales como en los sistemas agrícolas,
depende de la oferta ambiental y de nutrientes (Wensterman, 2005; Schilling et al., 2016),
que interactúan con los genotipos para producir la biomasa vegetal de la cual se obtienen
los productos cosechables para alimento (Zhao et al., 2017). Para el caso del cultivo de
papa, parte de esta energía requerida para la conversión de asimilados se usa en la
absorción, translocación y acumulación de nutrientes esenciales en el tubérculo (Sierra et
al., 2002; Villamil, 2005; Kumar et al., 2013), convirtiéndose en una de las especies de
mayor demanda nutricional por kilogramo de materia seca producida y con mayores
aportes nutricionales de los cultivos (White et al., 2009; Kumar et al., 2013) que sustentan
la seguridad alimentaria del mundo (Hardigan et al., 2017).
Los índices fisiológicos de crecimiento están afectados por factores del ambiente como la
intensidad de radiación, la temperatura y la disponibilidad de agua y nutrientes (Wolf et
al., 1990). En este sentido la alta disponibilidad de nitrógeno (N) aumenta el índice del
área foliar (IAF), la duración del área foliar (DAF) y la cobertura del suelo por el cultivo de
1. Critical Dilution Curve for N, P, and K and Leaf Area Index in Potato 3
papa, principalmente a través de una tasa más alta de aparición de hojas (ramificación) y
de expansión del área foliar (Vos y Biemond, 1992). La expansión del área foliar juega un
papel importante en la demanda principalmente de N y el aporte de fertilizantes de este
nutriente (Lemaire et al., 2007; Zhao et al., 2014; Chakwizira et al., 2016).
El N es el elemento esencial para la síntesis de proteínas, respiración, crecimiento de

tubérculo (Westermann, 2005; Kavvadias et al., 2012), en condiciones de suelos del
trópico frío, el N puede ser afectado por baja tasa de mineralización, bajas temperaturas,
suelos arcillosos y bajos contenidos de materia orgánica (Castro y Gómez, 2013),
aunque bajo excesos de N se puede presentar consumos de lujo y se reduce la
tuberización (Fandika, 2012; Ruza et al., 2013). Bajo deficiencia, se ha observado
reducción de la materia seca por disminución del área foliar y menor número de foliolos
que incide en una menor intercepción de luz y una menor tasa de fotosíntesis (Balemi et
al., 2009; Marouani et al., 2014).
Estudios realizados por Villamil (2005) con respecto al comportamiento de la

concentración de N en tejidos del peciolos y el estado fenológico de Capiro, mostraron
mayor asimilación de N en los primeros estados de crecimiento, posteriormente tiende a
disminuir con valores que pasaron de 6,6% alrededor de los 40 días después de siembra
(dds) a 3,3% hacia los 120 dds; la disminución se acentúa principalmente al iniciarse la
tuberización. Resultados similares fueron reportados por Sierra et al. (2002) para el cv.
Desiree en Chile, coincidiendo con una mayor acumulación de N en las etapas de
crecimiento de los primeros 75 dds.
El fósforo (P) promueve el crecimiento de las raíces (Aguilar-Acuña et al., 2006), la rápida
formación de tubérculos y su incidencia en la síntesis de almidón (Chung et al., 2014;
Leonel et al., 2016). En suelos fuertemente ácidos con pH menores a 5,5 de la zona
Andina, el P se fija y forma precipitados con hierro y aluminio lo cual disminuye su
disponibilidad (Hernández et al., 2012; Castro y Gómez, 2013). El P en Capiro en la
Sabana de Bogotá, presentó la mayor asimilación en la etapa de máximo crecimiento
alcanzando concentraciones de 0,8% en los peciolos alrededor de los 40 dds,
posteriormente se estabiliza a 0,5%. Dependiendo del rendimiento, un cultivo puede
extraer alrededor de 60 kg ha-1 de P2O5 (Villamil, 2005). En contraste Covarrubias et al.
(2005) encontraron para el cv. Alpha que la absorción de P es mayor al inicio de

tuberización (45 dds) y el máximo desarrollo del tubérculo.
El potasio (K) es esencial para la translocación de azúcares hacia los tubérculos,

regulación hídrica, respiración y síntesis de almidón, procesos fundamentales en el
crecimiento y llenado de tubérculos (Haeder et al., 1973; Perrenoud, 1993; Schilling et al.,
2016); la baja oferta edáfica en suelos arenosos, los excesos de Ca (Kavvadias et al.,
2012), la salinidad y la fijación por arcillas de tipo 2:1 limitan la disponibilidad de K en el
cultivo de papa (Castro y Gómez, 2013). El K es el elemento que más es extraido por
este cultivo. En Capiro ha presentado un 9% en el peciolo en el inicio de la floración o
cuando ha alcanzado el máximo índice de área foliar, cuando se presenta la mayor
asimilación.
Durante los primeros estadios de crecimiento y hacia el final del llenado del tubérculo, las
concentraciones foliares de K pueden alcanzar concentraciones foliares de 5,5%
(Villamil, 2005). Lo anterior difiere para los cultivares Desiree y Pimpernel en Chile,
donde el K se acumula en los primeros 70 dds y luego en la etapa de máximo
crecimiento del tubérculo entre los 100 a 130 dds con valores entre 8 y 10% en el peciolo
(Sierra et al., 2002), posiblemente por su condición de pertenecer a la subespecie
tuberosum y ser cultivada en zonas templadas bajo fotoperiodo de día largo.
Las relaciones alométricas son medidas indirecta del crecimiento, se utilizan para
pronosticar la concentración (Gilleto y Echevarria, 2015), consumo (Cogo et al., 2006),
eficiencia y pérdida de nutrientes en el ambiente (Abdallah et al., 2016). En el cultivo de
papa, las funciones alométricas se han utilizado principalmente con base en el peso seco
total para estimar las concentraciones críticas de nutrientes minerales, siendo N el más
común (Giletto y Echeverría, 2015; Abdallah et al., 2016) y con menos referencias P
(Zamuner et al., 2016) y K (Cogo et al., 2006).
La partición de nutrientes minerales en hojas, tallos y tubérculos durante el crecimiento

del cultivo de papa, pueden usarse para definir el estatus nutricional, fenómeno que
describe el comportamiento de la planta y explica las diferencias entre cultivares
mediante indicadores de partición de asimilados y nutrientes (Swain et al., 2014; Giletto y
Echeverría, 2015). La curva crítica de consumo de nutrientes es importante para
pronosticar la cantidad de nutrientes necesaria para producir una determinada biomasa

(Andriolo et al., 2006) de acuerdo al potencial de producción del cultivo.
Los conceptos de uso eficiente de nutrientes y fertilizantes generalmente describen el

adecuado comportamiento fisiológico en la planta en respuesta al uso de nutrientes
(Stewart, 2007; Prochnow et al., 2009). La eficiencia de absorción de los nutrientes se
estima por la diferencia de absorción del elemento entre plantas fertilizadas y no
fertilizadas, usando trazadores isotópicos, o medidas indirectamente a partir de la
acumulación de biomasa seca, que puede definir el concepto de eficiencia de utilización
del nutriente (EUN) y la eficiencia fisiológica del nutriente (EFN) (Dooberman, 2007;
Stewart, 2007; Prochnow et al., 2009), considerando factores de aportes y reciclaje
interno de nutrientes. El uso eficiente para P y K es menos estudiado en el cultivo de
papa (Fernandes y Soratto, 2012; Cogo et al., 2006).
El cultivo de papa es uno de los sistemas productivos con mayor consumo de nutrientes
y fertilizantes por ciclo y unidad de área, para Chile y Argentina estos índices pueden
oscilar entre 0,25 - 0,3 kg m-2 por semestre, respecto a cultivos como zanahoria y maíz
con 0,15 y 0,08 kg m-2 por semestre, respectivamente (Sierra et al., 2002). Un mayor
consumo de fertilizantes en papa, en estos países, se relaciona con altos índices de
cosecha y mayor potencial de rendimiento (50-60 t ha-1) obtenidos por la tecnificación del
sistema especialmente para especies de la subespecie tuberosum cvs. Pimpernel y
Desireé (Sierra et al., 2002; Aguilar-Acuña et al., 2006).
En Colombia se han realizado estudios importantes respecto a la fisiología de

crecimiento y desarrollo para los cultivares de papa como Criolla Colombia (Valbuena et
al., 2010; Santos, 2010), Capiro y Suprema (Santos et al., 2010), con un incremento en el
potencial productivo en Capiro (50-80 t ha-1) y Suprema (40-70 t ha-1), las dos del grupo
Andigena, debido a un manejo adecuado de semilla certificada, riego, fertilización
específica y mecanizada con el índice de consumo de fertilizantes de 0,2-0,3 kg m-2
(Villamil, 2005; Ríos et al., 2010; Gómez y Torres, 2012), pero no han relacionado ni
cuantificado la acumulación y distribución de nutrientes.
La absorción de nutrimentos durante las etapas fenológicas en cultivares comerciales de

papa del grupo Andígena han sido poco estudiados en Colombia, donde los últimos
estudios detallados se realizaron en la década de los 70 para los cultivares ICA Puracé y
Guantivá (Grandet y Lora, 1979).
En los últimos años, trabajos realizados por Villamil (2005) y Gómez y Torres (2012) han
permitido conocer de manera general los niveles foliares y aproximaciones a los
requerimientos nutricionales en Capiro, pero sin una relación de su acumulación y
distribución bajo diferentes ofertas edáfico-ambientales y de aportes nutricionales
variables. La información disponible de fertilización y manejo de nutrición está
encaminada al manejo de respuestas en campo de N, P y K, pero no hay una correlación
con las demandas nutricionales de cada cultivar, ciclo de desarrollo, potencial de
rendimiento e incluso destino de producción (Villamil, 2005).
Dentro de los costos de producción, el manejo de la fertilización incluye el uso de

fertilizantes edáficos, foliares y enmiendas orgánicas que pueden oscilar entre el 22 y
35% del costo total, con un valor equivalente entre 5 y 6 millones de pesos/ha (Gómez y
Torres, 2012; Pérez et al., 2014). En Colombia, los insumos para la fertilización del
cultivo, ocupan un lugar destacado en los costos totales de producción de papa. En
Capiro y Suprema, la inversión en fertilización alcanza en promedio el 23% de los costos
totales. La variación en el rubro y manejo de la fertilización-enmiendas depende del
sistema tecnológico propio dado por el agricultor (mecanización, riego, semilla certificada,
gestión de los procesos agronómicos, técnicas de fertilización) (Gómez y Torres, 2012;
Pérez et al., 2014).
La investigación en nutrición de papa en Colombia se limita a respuestas en dosis y

fuentes y no se han actualizado requerimientos nutricionales ni indicadores de
diagnóstico nutricional (Villamil, 2005; Cevipapa, 2005). Además, no se realiza un manejo
específico de los nutrientes de acuerdo al tipo de suelo, cultivar, fenología en el altiplano
Cundi-Boyacense que afecta el potencial de rendimiento (Gómez y Torres, 2013).
Adicionalmente, un desconocimiento en la distribución y partición de nutrientes y su
relación con índices de crecimiento y uso eficiente de nutrientes.
Con el fin de abordar la problemática anterior, se establecieron las siguientes hipótesis:

(i) no existe diferencias en el crecimiento con el uso de fertilización variable y ciclos del
cultivo para los cultivares Diacol Capiro y Pastusa Suprema; (ii) la extracción de N, P y K
en los tubérculos dependen del genotipo y no del factor ambiente (suelo) y la fertilización
aplicada y (iii) la partición de N, P y K no necesariamente concuerda con la partición de
biomasa debido a las caracteristicas morfológicas del cultivar y a la funcionalidad
diferencial de N, P y K.
Para responder a las anteriores hipótesis se plantearon los siguientes objetivos:
Objetivo general
Caracterizar la acumulación y distribución de macronutrientes en dos cultivares de

Solanum tuberosum L. en diferentes ambientes en el altiplano Cundi-Boyacense.
Objetivos específicos
Caracterizar la distribución de macronutrientes mediante la partición diferencial de

elementos minerales en hojas, tallos y tubérculos según ambiente de evaluación,
cultivar y niveles de fertilización.
Determinar la acumulación de macronutrientes minerales en las diferentes etapas

fenológicas por cultivar, niveles de fertilización y el efecto de la oferta edáfica-
ambiental en cada ambiente de evaluación.
Establecer la relación entre la acumulación de nutrientes minerales, variables

fisiológicas de crecimiento y uso eficiente de nutrientes por cultivar.
El presente trabajo está elaborado como un compendio de cinco artículos que

corresponde a cada capítulo en los cuales se desarrollan los diferentes aspectos
requeridos para el cumplimiento de los objetivos:
Capítulo 1. Critical Dilution Curve for N, P, and K and Leaf Area Index in Potato (Solanum
tuberosum L., Group Andigena) in Colombia. Sometido en Agronomy Journal
Capítulo 2. Uptake and partition of N, P, and K in potato (Solanum tuberosum L., Group
Andigena). Sometido en Field Crops Research Journal.
Capítulo 3. Accumulation of N, P, and K in the tubers of potato (Solanum tuberosum L.)

spp. andigena under contrasting soils of the Andean region of Colombia. Publicado en
Agronomía Colombiana 35(1), 56-67, 2017. Doi: 10.15446/agron.colomb.v35n1.61068
Capítulo 4. Potential yield and efficiency of N and K uptake in tubers of cvs. Capiro and
Suprema (Solanum tuberosum subsp. andigena). Sometido en Agronomía Colombiana.
Capítulo 5. Diagnóstico de K+ y NO3- en savia para determinar el estado nutricional en

papa (Solanum tuberosum L. subsp. andigena). Publicado en la Revista Colombiana de
Ciencias Hortícolas 11(1), 133-142, 2017. Doi: 10.17584/rcch.2017v11i1.6132
Los capítulos 1, 2, 3 y 4 fueron proyectados para su publicación en revistas de habla

inglesa, de tal manera que se elaboraron en dicho idioma. El capitulo 5 y las secciones
restantes fueron elaborados en español. Dado que el trabajo se presenta como un
compendio de artículos, existen algunas diferencias menores de estilo, relacionadas
principalmente con formatos de citas bibliográficas en el texto y elaboración de
referencias bibliográficas.
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1. Critical Dilution Curve for N, P, and K and
Leaf Area Index in Potato (Solanum
tuberosum L., Group Andigena)1
Curvas de dilución crítica para N, P y K e índice de área

foliar en papa (Solanum tuberosum subsp. andigena en
Colombia)
Manuel Iván Gómez S., Stanislav V. Magnitskiy, Luis Ernesto

Rodríguez
1.1 Abstract
The critical dilution curves (CDC) for nitrogen (Nc), phosphorus (Pc), and potassium (Kc)
obtained from total dry biomass (W) and leaf area index (LAI) were used as a diagnostic
tool to determine nutrient status and to adjust fertilization rates in two cultivars of potato
Group Andigena. The objective of this research was to determine allometric ratios of Nc,
Pc, and Kc (%) and nutrition index (NI) based on W and LAI in cultivars (cv.) Diacol
Capiro (Capiro) and Pastusa Suprema (Suprema). Additionally, optimal fertilization rate
was evaluated to achieve maximum yields during two growth cycles in contrasting
environments of the Andean zone of Colombia. The CDC was validated using W in cv.
–0.3197 –0.198 –0.269
Capiro: Nc = 6.23W ; Pc = 0.523W ; Kc = 9.02W and in cv. Suprema: Nc =
–0.327 –0.186 –0.1353
6.74 W ; Pc = 0.536W ; Kc = 6.585W and presented higher robustness
than the CDC obtained from LAI. Cv. Capiro presented a lower dilution coefficient b than
cv. Suprema due to the higher physiological efficiency in tuber growth. The NI ranged
between 0.25-1.32 with better fit in cv. Capiro and relative yields (RY) starting from 40%
1
Artículo sometido en Agronomic Journal
without fertilization; for cv. Suprema, null to marginal response to fertilization was
obtained indicating a luxury uptake of N (NI 1-1.5) with RY starting from 70%. Except for
this research, Nc, Pc, and Kc have not been validated for cultivars of Group Andigena.
The CDC for this cultivated Group could serve to identify deficiency, sufficiency, or excess
of N, P, and K and to predict final yield per cultivar under highland equatorial conditions.
Key words: nutrition index, allometric relationships, dry biomass, Solanaceae.
Core Ideas
• Diagnostics of mineral nutrition in field is poorly developed for potato Group Andigena.
• Critical dilution curves CDC for N, P, and K were validated in potato Group Andigena.
• The CDC serve to identify macronutrient status and to predict yield of potato in the
tropics.
Abbreviations: CDC, critical dilution curve; W, total plant biomass; LAI, leaf area index;
Nc, total critical concentration of mineral nutrient; NI, nutrition index; RY, relative yield;
ODF, optimal dose of fertilizer; MY, maximum yield.
1.2 Resumen
Las curvas de dilución crítica (CDC) para nitrógeno (Nc), fósforo (Pc) y potasio (Kc)
estimadas a partir de la biomasa seca total (W) y el índice de área foliar (IAF) se usan
como métodos de diagnóstico, para determinar el estado nutricional y realizar ajustes en
la fertilización para optimizar el potencial de rendimiento en papa. El objetivo de este
estudio fue determinar las CDC mediante relaciones alométricas de Nc, Pc y Kc (%) y el
índice de nutrición (INN) con base en W y el IAF en los cultivares Diacol Capiro (Capiro) y
Pastusa Suprema (Suprema). Adicionalmente, se determinó la dosis óptima de
fertilización para obtener mayores rendimientos en dos ciclos de cultivo en ambientes
contrastantes de la zona Andina de Colombia. Se determinó la CDC a partir de W
encontrando para Capiro: Nc = 6,23 W-0,319; Pc = 0,523W -0,19
; Kc = 9,02W -0,269
, mientras
-0,327 -0,186 -0,1353
para Suprema: Nc= 6,74 W ; Pc= 0,536W ; Kc= 6,585W , observandose
mejor robustez comparado con IAF. Capiro presentó un menor coeficiente de dilución b
que Suprema por su mayor eficiencia fisiológica en el crecimiento de tubérculos. El INN
presentó rangos entre 0,25-1,32 con mejor ajuste en Capiro y rendimientos relativos (RR)
desde 40% sin fertilización, para Suprema se observó respuesta nula a marginal en la
fertilización mostrando consumo de lujo de N (INN 1-1,5) con RR desde 70%. Las CDC
pueden identificar situaciones de deficiencia, suficiencia y exceso de N, P y K y puede
ser usadas para predecir el rendimiento final por cultivar.
Palabras clave: Indice de nutrición, relaciones alométricas, biomasa seca, Solanaceae.
1.3 Introduction
Potato (Solanum tuberosum L.) is one of the species with the highest nutritional demand
in nitrogen (N), phosphorus (P), and potassium (K) per kg of dry matter produced (Kumar
et al., 2013, Gómez et al., 2017a). In the Colombian Andean region, potato cultivation
requires a high consumption of fertilizers per cycle and unit area (0.2-0.3 kg m-2 of
fertilizer) to obtain high yields and product quality (Gómez and Torres, 2012). However,
the optimal fertilization dose of N (Giletto and Echeverría, 2015), P (Fernandes and
Soratto, 2016) and K (Zelelew et al., 2016) can vary widely between cultivars and per site,
so that a proper management of these mineral nutrients favors the profitability and
sustainability of potato productive system.
Diagnostics methods in plants can be used to improve nutrient efficiency and reduce
nutrient losses in the environment (Abdallah et al., 2016). The critical dilution curves
(CDC) are established between the concentrations of N, P and K (%) in organs in relation
to the dry biomass (W) (Chakwizira et al., 2016, Abdallah et al., 2016), leaf area index
(LAI) (Ata-Ul-Karim et al., 2014; Zhao et al., 2014) by means of direct measurements in
plants or indirect estimation, such as through a duration of leaf area (Wang et al., 2017).
Expansion of leaf area plays an important role in the main demand of N and contribution
of N fertilizers (Lemaire et al., 2007; Zhao et al., 2014; Chakwizira et al., 2016) and is
considered a fundamental growth variable to monitor environmental and agronomic
variations due to the capacity of light interception by canopi and crop photosynthetic
capacity (Ata-Ul-Karim et al., 2014). In potato crop, allometric equations have been based
mainly on total dry weight to estimate critical concentration of mineral nutrients, such as N
that is more common (Giletto and Echeverria, 2015; Abdallah et al., 2016) and, with fewer
references, P (Zamuner et al., 2016) and K (Cogo et al., 2014).
The critical value of the mineral nutrient is defined as a minimum concentration of nutrient
needed for the crop to reach maximum growth rates at given biomass accumulation
(Greenwood et al., 1990; Lemaire et al., 2007; Raut, 2017) (Equation 1) or at given LAI
(Ata-Ul-Karim et al., 2014; Wang et al., 2017) (Equation 2). The (CDC) for nitrogen (Nc)
could be represented by a negative exponential model:
Nc = aW-b (1)
Nc = aIAF-b (2),
where W is a total biomass (t ha-1), the LAI is a leaf area index, Nc (%) is a total critical
concentration of the nutrient (N, P or K) in total biomass (g 100 g -1), the coefficient a
represents the concentration of nutrient when total biomass is lower than 1 t ha -1, and the
parameter b represents a "dilution coefficient", which describes the reduction in the
concentration of the nutrient to the extent that total biomass increases.
CDC of mineral nutrients nutrient is used to diagnose nutrient deficiency (Bélanger et al.,
2015), adjust fertilization (Gilleto and Echevarria, 2015; Zamuner et al., 2016), or simulate
demand of the nutrient and yield in productive systems (Lemaire et al., 2007). Thus, for N,
Nc defines three levels of this mineral element in plants (Marouani et al., 2014): (i) values
significantly below the curve represent crop growth limited by the supply of the nutrient,
(ii) values above the curve represent the growth under a luxury consumption of nutrient
(iii) the values on the curve represent the growth in the Nc, in other words, the adequate
level of nitrogen for an optimum crop growth.
In potato, the values of parameters a and b have been estimated using the sum of
biomass of leaves, stems, and tubers with respect to the concentration of mineral
nutrients in these organs under contrasting edapho-climatic conditions with different
cultivars and under different rates of nutrients (Marouani et al., 2014, Zamuner et al.,
2016). In other studies for ssp. tuberosum, Greenwood et al. (1990) reported values 5.36
and -0.46 for coefficients a and b, respectively, while Gilletto and Echevarria (2015)
obtained values for a ranging from 5.19 to 5.53 and for b ranging from -0.25 to -0.42 in
Russet cultivars in Argentina.
Evaluation of CDC based on dry biomass is more common than the one obtained by LAI
(Wang et al., 2017). Evaluation for Nc that for Pc is more usual for ssp. tuberosum
(Abdallah et al., 2016; Zamuner et al., 2016) and currently there are no reports for Nc, Pc
and Kc in ssp. andigena. Andriolo et al. (2006) in Brazil evaluated Nc in cv. Axterix, while
the other studies were done by Marouani et al. (2014) in Tunisia in cvs. Spunta and Bellini
and Gilleto and Echeverria (2015) in cvs. Gen Russet, Umatilla Russet, Bannock Russet,
and Markies Russet in Argentina. On the other hand, CDC evaluation of Pc is less
common and was initially established by Zamuner et al. (2016) for ssp. tuberosum in cv.
Innovator in Argentina. Currently, there are no reports on evaluation of Kc in potato in the
field but only in greenhouse conditions for cv. Asterix by Cogo et al. (2006). Our study
might be one of the first CDC references regarding LAI for N, P and K in potato ssp.
andigena; although it has been evaluated in other crops, such as rice (Ata-Ul-Karim et al.,
2014) and wheat (Zhao et al., 2014; Wang et al., 2017).
The objectives of this study were (i) to establish allometric relationships for Nc, Pc, and Kc
(%) and NI based on the total dry biomass (W) and leaf area index (LAI) in cvs. Capiro
and Suprema; (ii) compare CDC evaluated in ssp. andigena with that evaluated in ssp.
tuberosum; (iii) determine the optimal fertilization dose to achieve the maximum yields in
cvs. Capiro and Suprema. This study was undertaken under variable fertilization rates at
five stages of growth and during two growth cycles in soils that were contrasting in fertility.
The research was done in 2013-2016 in the Andean region of Colombia to provide a
diagnostics and timely management of N, P, K fertilizers and to predict the crop yield.
1.4 Materials and methods
1.4.1 Field experiments

The field evaluation was carried out in two production cycles (2013-2016) in two locations
of altiplano Cundiboyacense (Colombia): Facatativá (soils of high fertility saturated in
bases, Andic Eutrudepts) and Chocontá (desaturated acid soils of low fertility, Humic
Dystrudepts). These locations were representative for the region and presented high yield
potential and contrasting edaphoclimatic conditions (Table 1-1).
Table 1-1. Environmental and soil fertility characteristics in studied locations.
Facatativá Chocontá
Environmental conditions††
Cycle 1 Cycle 2 Cycle 1 Cycle 2
/ Location
(I-2013) (I-2015) (II-2013) (I-2016)
Altitude, m.a.s.l. 2597 2597 2780 2710
North Latitude 4°49´26.9” 4°49´39.9” 5°5´30.37” 5°6´23.94”
West Longitude 74°22´29.7” 74°22´49.3” 73°43´2.04” 73°40´48.53”
Annual rainfall, mm 951 850 1295 1058
Rainfall per crop cycle, mm 397 415 712 803
ET per crop cycle, mm 454 382 640 603
Max air temperature, ºC 18.1 18.5 16.2 16.5
Min air temperature, ºC 7.0 7.2 4.4 10.1
Average air temperature, ºC 12.7 12.5 10.6 12.9
Soil Andic Andic Humic Humic
properties/classification† Eutrudepts Eutrudepts Dystrudepts Dystrudepts
Texture Loam Loam Clay Loam Clay Loam
Soil fertility††† High High Low Low
pH 6.4 5.8 5.5 5.3
–1
Al, cmolc kg 0 < 0.1 0.1 0.54
–1
Organic matter, g kg 166.7 127.1 67.7 85.9
–1
CEC, cmolc kg 31.95 19.14 9.52 7.9
–1
N, g kg 8.3 6.4 3.3 4.3
–1
P, mg kg 39.64 70.16 18.18 41.5
–1
K, cmolc kg 3.14 0.87 0.68 0.84
†Physical-chemical analysis of soil in arable layer (0-30 cm) was done according to IGAC (2006).
Classification of soils followed USDA classification system (Soil Survey Staff, 2014).
††Environmental conditions data were taken and calculated from IDEAM (2017). †††Potential
chemical fertility was evaluated in arable layer 0-30 cm of soil (Castro and Gómez, 2013).
For each location and cycle, the experiment was established using incomplete factorial
design in divided subplots – a mixed model with four replicates in complete blocks at
random, where the main plot corresponded to cultivars (Capiro or Suprema) and the
subplots corresponded to four levels of fertilization (F0, F1, F2, or F3), where F0 were
unfertilized plots (initial conditions of soil fertility). The contribution of mineral nutrients
from fertilizer treatments to each location and in each cycle, the sources of fertilizers and
dose fractionation are presented in Table 1-2.
Table 1-2. Doses of mineral nutrients in fertilizer treatments.

†Nutrient Cycle 1 Cycle 2
Location dose (kg F1 F2 F3 F1 F2 F3
–1
ha ) 1186 1582 1977 1450 1900 2375
N 128 171 214 123 164 205
P2O5 196 261 326 216 288 360
K2O 135 180 225 176 235 294
Mg 53 70 88 60 80 100
Facatativá S 56 74 93 113 150 188
B 2.55 3.4 4.3 1.7 2.3 2.9
Zn 4.2 5.6 7.0 3.5 4.6 5.8
Mn 5.25 7.0 8.8 4.2 5.6 7.0
Total 1632 2175 2719 1500 2000 2500
N 144 192 240 143 191 239
P2O5 255 340 425 285 380 475
K2O 261 348 435 236 315 394
Mg 42 56 70 30 40 50
Chocontá
S 90 120 150 29 38 48
B 0.9 1.2 1.5 3.3 4.4 5.5
Zn 1.8 2.4 3.0 3.6 4.8 6.0
Mn 2.3 3.0 3.8 4.1 5.4 6.8
†Fertilization rates were recommended by soil-plant balance method (Castro and Gómez, 2013)
and fractionated according to historical practices carried out in the areas, where high yields (> 50
Mg ha–1 tuber fresh weight) were obtained: N, 60% at planting and 40% at 45-50 dap; P, 70% at
planting and 30% at 45-50 dap; K, 30% at planting and 70% at 45-50 dap. Granulated fertilizers
were used: Diammonium phosphate (18-46-0) for N and P; K, KCl (0-0-60) and Potassium Sulfate
®
(0-0-50); Ca, Calcium Nitrate (25% CaO); Mg, Kieserite; Nutricomplet – Ingeplant Colombia
(complex source of B, Zn, Cu, Mn and S, based on sulfates).
Thirty two combinations of factors were evaluated using repeated measures design with
four factors: two growth cycles (I, II); two cultivars (Capiro, Suprema); two locations
(Facatativá and Chocontá) and four levels of balanced fertilization (Table 1-2), with an
intra-subject factor of time associated with five critical phenological stages of crop growth
(Valbuena et al., 2010): Stage I, 50-55 days after planting (dap) (formation of branches
and primary stems); Stage II, 70-75 dap (formation of secondary stems and initiation of
tuberization); Stage III, 90-100 dap (flowering, maximum tuberization, and start of tuber
filling); Stage IV, 120-125 (end of flowering, filling of tubers); Stage V, 150-160 dap
(senescence, maximum filling, and maturation of tubers). Planting was done in
experimental units of 50 m2 (135 plants plot–1), with 1 m between the rows and 0.37 m
between the plants, a useful area of 36 m 2 (three central rows) and a density of 27000
plants ha-1. Irrigation was applied following indications of weather stations (IDEAM, 2017)
and weed and phytosanitary controls were carried out according to the needs of the crop.
1.4.2 Analytical methods

Four plants were selected per experimental unit and growth stage and a destructive
analysis of leaves, stems, and tubers was carried out. At each sampling moment, each
organ was weighed in fresh for each plant separately. For chemical analysis, all plant
parts were rinsed with deionized water, then the same organs from four plants were
mixed to receive a subsample of 200 g (fresh weight) that was dried in an oven at 70°C
for 72 h and the dry weight (DW) was measured (IGAC, 2006).
All organs were subjected to chemical analysis according to IGAC (2006). Total contents
of N, P and K in each organ were estimated by multiplying nutrient concentrations in the
organ (g 100 g–1 DW) by the amount of dry biomass accumulated per stage (Mg ha –1)
according to Abdallah et al. (2016). Leaf area (cm 2) was determined in green plants using
a LI-COR leaf area meter (LI-3100, Lincoln, NE, USA) and LAI was estimated (Ata Ul
Karim et al., 2014). In both cycles and locations, the yield at Stage V was evaluated at
150-160 dap (senescence, maximum filling, and tuber maturation).
1.4.3 Data analysis

Nutrient dilution curves for Nc, Pc and Kc in relation to total dry mass W (plant biomass
without roots and stolons) (Mg ha –1) were elaborated according to equations (1) and (2),
with the procedure adapted from Greenwood et al. (1990) for potato, Giletto and
Echeverría (2015) for Nc, Zamuner et al. (2016) for Pc; the previous models for Kc were
adjusted since no current references were found for this mineral element. On the other
hand, the procedure of Ata-Ul-Karim et al. (2014) was adapted to obtain CDC based on
LAI. The calibration of critical curves required identification of values by which the
evaluated nutrients might not limit crop growth (Abdallah et al., 2016; Wang et al., 2017).
For this reason, analysis of variance was done by cultivar, location, and crop cycle at
each sampling stage with respect to W and LAI and the total concentrations of nutrients
(N, P, and K) in plants (leaves, stems, and tubers). When significant differences were
found between fertilizer doses (Table 1-2), the data was selected with concentration of
nutrient that expressed maximum accumulation of DW or LAI using LSD test (P < 0.05).
From the selected data, a non-linear regression was performed to adjust the data to
allometric curve. The coefficient of determination (R 2) was calculated from the sum-
squared relation of error and total (adjusted coefficient), where 95% confidence intervals
were constructed for each of the estimators to evaluate differences of models among the
cultivars.
The nutrition index (NI) and relative yield (RY) that characterized nutritional status of
plants were calculated. For each cultivar, the NI for N (i.e. NNI) was obtained by dividing
the N concentration (%) in W by the Nc at each sampling date in different locations and
growth cycles (Bélanger et al., 2001; Giletto and Echeverría, 2012). The RY was
calculated as a ratio of the harvested yield (tuber fresh weight, Mg ha –1) obtained at each
fertilization rate to the highest yield (tuber fresh weight, Mg ha –1), multiplied by 100
(Marouani et al., 2014). The optimum dose of fertilizer (ODF, kg ha -1), which provides the
maximum yield (MY, Mg ha–1), was obtained from the first derivative of a quadratic
polynomial model between the fertilizer dose and yield for each cultivar, cycle, and
location according to Giletto and Echeverría (2015). This model defines the ODF as
follows: Y = a + bX + cX2, where Y corresponds to RY, X is fertilizer dose (kg ha -1), c is
quadratic coefficient, b is linear coefficient, and a is intercept; therefore, with dy/dx = 0,
the ODF is equal to c/2b (Giletto and Echeverría, 2015). In the present study, the RY was
expressed as a function of NI and the difference of the optimal fertilizer dose (dODF)
under a quadratic model was adapted from Giletto and Echeverría (2015).
For variables yield (tuber fresh weight, Mg ha –1) and LAI evaluated at 150-160 dap, a
mixed model was proposed assuming the replicates were random effects, while the main
effects, cultivation, location, crop cycle, and fertilizer levels were assumed as fixed effects
considering all the possible interactions between the main effects. After that analysis of
variance mean comparisons was made for each pair of combinations Cultivar x Location x
Cycle x Fertilization using LSD test (P < 0.05). For all statistical analyzes described, SAS
9.4 software was used (SAS Institute, 2014).
1.5 Results and discussion
1.5.1 Critical Dilution Curves for Nc, Pc, and Kc
The critical curves for Nc, Pc, and Kc in both cultivars followed a negative exponential
model in relation to W (Figure 1-1) and LAI (Figure 1-2). According to the curves, the
concentration of mineral nutrients decreased during expansive growth (AF) and growth in
mass (W); the total biomass increased more at stages of tuber filling because of a higher
accumulation of assimilates in sink organs complying with a "dilution law" proposed by
Justes et al. (1994). This model in cultivars of Group Andigena was similar to the ones
described in Group Chilotanum by Greenwood et al. (1990) for N, Zamuner et al. (2016)
for P, and Cogo et al. (2006) for K based on W and by Ata-Ul-Karim et al. (2014) and
Wang et al. (2017) for C3 plants based on LAI.
For Nc, no significant differences were observed between the cultivars in coefficients a
and b based on W, with coefficient a ranging from 6.23 to 6.74 and dilution coefficient b
ranging from -0.3197 to -0.327 for cvs. Capiro and Suprema, respectively (Table 1-3);
these coefficients were higher than those reported in Group Chilotanum by Greenwood et
al. (1990); Andriolo et al. (2006); Giletto and Echeverría (2012); Giletto and Echeverría
(2015) in cv. Gem Russet; Marouani et al. (2014) in cv. Bellini and by Abdallah et al.
(2016) in cv. Bintje. The coefficients b obtained in the present study were inferior to those
obtained by Giletto and Echeverría (2015) in cv. Markies (-0.250) and similar to ones
obtained in cv. Spunta (-0.310) by Marouani et al. (2014) (Table 1-3). All references used
for the comparison (Table 1-3) referred to total dry biomass in the same terms as the
present study employing W as a sum of above-ground and tuber matter.
10 Capiro
N concentration Nc (%)
9
8 Suprema
7
6
5
4
3
2
1
0
(a) 0 5 10 15 20 25 30
0.8
0.7
0.6
P concentration Pc (%)
0.5
0.4
0.3
0.2
0.1
(b) 0
0 5 10 15 20 25 30
12
10
K concentration Kc (%)
0
0 5 10 15 20 25 30
(c)
Total dry biomass (Mg ha-1)
Figure 1-1. Critical dilution curves for N (A), P (B), and K (C) in cvs. Capiro ( ) and
Suprema ( ) in two production cycles (2013-2016) under non-limiting conditions of
fertilization in soils of contrasting fertility. Means consisted of n = 13 data per phenological
stage and cultivar. Coefficients a and b of potential allometric function were based on total
dry biomass W (equation 1).
8 Capiro
7
Suprema
N concentration Nc (%)
6
5
4
3
2
1
0
(a) 0 2 4 6 8 10
0.6
0.5
P concentration Pc (%)
0.4
0.3
0.2
0.1
(b) 0
0 2 4 6 8 10
10
9
8
K concentration Kc (%)
7
6
5
4
3
2
1
0
(c) 0 2 4 6 8 10
Leaf area index (LAI)
Figure 1-2. Critical dilution curves for N (A), P (B) and K (C) based on LAI in cvs. Capiro
( ) and Suprema ( ) in two production cycles (2013-2016) under non-limiting conditions
of fertilization in soils of contrasting fertility. n = 65 data per cultivar. Coefficients a and b
of allometric function were based on LAI (equation 2).
Table 1-3. Allometric relationships and critical dilution coefficients for N, P, and K based
on total dry biomass W (above-ground organs and tubers) for cvs. Capiro and Suprema
as compared with cultivars of Group Chilotanum. The data for cvs. Capiro and Suprema
were obtained under non-limiting conditions of fertilization in soils of contrasting fertility.
Dilution curve Interval of Interval of

Range W 2
Nutrient Cultivar –1 R confidence confidence SEa SEb
(Mg ha )
–b a 95% -b 95%
Nc = aW
Capiro 6.23 W –0.3197 1-22 0.93 5.55-6.90ns 0.26-0.38ns 0.3375 0.0318
–0.327 ns ns
Suprema 6.74 W 1-24 0.91 5.83-7.65 0.24-0.39 0.4578 0.0377
1 –0.420
Innovator 5.30 W 1-14 0.92 - - - -
2 –0.580
Gem Russet 5.32 W 1-19 0.76 - - - -
Markies2 5.53 W –0.250 1-24 0.89 - - - -

N Bintje3 5.37 W –0.45 1-12.7 0.86 - - - -
cv. Group
4
5.36 W –0.460 1-15 - - - - -
Chilotanum
5 –0.310
Spunta 3.25 W - 0.99 - - - -
Bellini5 2.99 W –0.380 - 0.94 - - - -
Asterix6 3.60 W –0.370 1-13 0.88 - - - -

–0.198 ns ns
Capiro 0.523 W 1-22 0.95 0.47-0.57 0.15-0.25 0.0256 0.0252
–0.186 ns ns
P Suprema 0.536 W 1-24 0.95 0.45-0.63 0.11-0.26 0.0444 0.0392
Innovator7 0.392 W –0.304 1-16 0.91 - - - -
Capiro 9.02 W –0.269 1-22 0.91 7.78-10.3** 0.19-0.34** 0.6205 0.0400
K Suprema 6.585 W –0.1353 1-24 0.92 5.38-7.76** 0.05-0.18** 0.600 0.0399

8 –0.317
Asterix 5.54 W 1-7 0.90 - - - -
W, total dry biomass; a and b, coefficients of critical dilution curve (NC) for N, P, and K under non-
limiting conditions of fertilization. SE, standard error of means of coefficients. **Significant
differences (P<0.05) of coefficients a and b between the studied cultivars; ns, not significant.
1
Giletto and Echeverría (2012); 2Giletto and Echeverría (2015); 3Abdallah et al. (2016);
4
Greenwood et al. (1990); 5Marouani et al. (2014); 6Andriolo et al. (2006); 7Zamunner et al. (2016);
8
Cogo et al. (2006).
As for Pc for cvs. Capiro and Suprema, coefficient a ranged within 0.523 and 0.536 and
dilution coefficient b varied from -0.198 to -0.186, respectively; the coefficient b did not
present differences among the cultivars being higher in the cultivars of Group Andigena
than those reported for cv. Innovator by Zamuner et al. (2016) (Figure 1-1B and Table
1-3). Similar ranges for Pc were reported in wheat by Bélanger et al. (2015) and Raut et
al. (2017). The highest coefficients a for Nc and Pc in the studied cultivars could be
associated with higher requirement of N and P at the initial stages of growth in this crop,
such as required for stolon branching and differentiation of tubers that characterize these
cultivars corroborating data by Santos et al. (2010).
The highest coefficients b for Pc could indicate a lower efficiency in assimilation and
translocation of N and P in cvs. Capiro and Suprema at tuber filling due to their higher
production of W up to 30 to 35 Mg ha –1 as compared to Group Chilotanum with W of 24
Mg ha–1 according to Giletto and Echeverría (2015). The low efficiency of Pc in cultivars of
Group Andigena could be associated with fixation of P in soils of Andean region that
requires application of higher doses of P 2O5 (220 to 350 kg ha–1) with respect to
applications of 50-130 kg ha–1 P2O5 in cultivars of Group Chilotanum in temperate regions
of the world (Zamuner et al., 2016).
The coefficients a and b for Nc and Pc did not differ among the cultivars, possibly
because of the accumulation of these nutrients mostly in the aerial part with low
dependency on tuber growth. These results suggest a higher demand for N and P in
cultivars of Group Andigena as compared to Group Chilotanum to accumulate the same
total biomass. This was due to the higher dry weight of total biomass W that reached cvs.
Capiro (22 Mg ha–1) and Suprema (24 Mg ha –1) for being plants of average to high height
as compared to smaller plants with lesser dry biomass in Group Chilotanum (Table 1-3).
This favors higher requirements for N and P in cvs. Capiro and Suprema necessary for
continuous formation of organs (leaves, stems, and tubers), since these cultivars have a
longer vegetative period and are plants of greater size under conditions of the cold
tropics.
In contrast, intra-specific variability with significant differences between the cultivars was
observed for Kc in relation to coefficient a with respect to W (Table 1-3) and LAI (Table
1-4); a higher K requirement with 27% increase in K concentration was obtained in cv.
Capiro than in cv. Suprema at initial stages of growth (Stage I, formation of branches and
primary stems). At this stage Kc in cv. Capiro ranged between 7.5 and 10% K and
coefficient a was 9.02 for W and 8.84 for LAI, while cv. Suprema presented lesser Kc of 5-
7.4% and coefficients a of 6.58 (for W) and of 5.59 (for LAI); the lower values of Kc (5.56)
were reported for Group Chilotanum by Cogo et al. (2006) in cv. Asterix under
greenhouse conditions. The previous findings corroborate existence of genotypic
differences between cultivars of Group Andigena in demand for K despite similar
accumulation of dry biomass (Figure 1-1 and Figure 1-2 C). This was due to the lower
coefficient a for Kc in cv. Suprema at early stages of growth, which was related to higher
vegetative growth with LAI of 0.9-3.1 as compared to cv. Capiro with LAI of 0.5-2.1 (Table
1-4), thus, favoring “dilution” of K in cv. Suprema since this mineral element is not part of
the organic compounds in the plant cells. The phenomenon of dilution by expansive
growth was also explained by Lemair et al. (2007) and Chakwizira et al. (2016).
Table 1-4. Allometric relations and critical dilution coefficients for N, P, and K based on
leaf area index (LAI) for cvs. Capiro and Suprema under non-limiting conditions of
fertilization in soils of contrasting fertility.
Interval of Interval of
Dilution curve Range 2
Nutrient Cultivar R confidence (a) confidence SEa SEb
of LAI
NC = aLAI–b 95% (-b) 95%
–0.235 ns
N Capiro 4.33 LAI 0.5-6.3 0.84 3.57-5.09 0.08-0.39ns 0.3808 0.0762
–0.250 ns ns
Suprema 4.90 LAI 0.9-7.1 0.83 3.77-6.02 0.06-0.44 0.5618 0.0930
–0.082 ns ns
P Capiro 0.39 LAI 0.5-6.3 0.91 0.33-0.45 0.04-0.21 0.0293 0.0618
–0.090 ns ns
Suprema 0.41 LAI 0.9-7.1 0.88 0.31-0.51 0.08-0.26 0.0490 0.0861
–0.437 ns
K Capiro 8.84 LAI 0.5-6.3 0.89 7.41-10.3** 0.28-0.59 0.7132 0.0768
–0.149 ns
Suprema 5.95 LAI 0.9-7.1 0.91 4.79-7.10** 0.05-0.30 0.5787 0.0729
SE, standard error of coefficients; IC, Interval of confidence 95%. ** Significant differences p <0.05
of coefficients a and b between the study cultivars; ns, non-significant differences.
On the other hand, for Kc in cv. Capiro, a lower dilution coefficient b was observed at both
allometric ratios with significant differences (P < 0.05) as compared to cv. Suprema
(Table 1-3 and Table 1-4). The Kc concentrations at maturity stage were equal to 3.5% in
cv. Capiro and 4.2% in cv. Suprema with 30 Mg ha –1 W obtained (Figure 1-1C), which
may explain a more efficient use of K in cv. Capiro since it presented a better
translocation than cv. Suprema at both locations and cycles with higher production of
tubers and biomass per kg of K uptake (data not shown). Regarding dilution coefficient b
Cogo et al. (2006) obtained the lower values in a greenhouse study (-0.317). This
research corroborated the existence of a "dilution phenomenon” for K considering that this
element does not enter into a structural pool of the cells but locates in the metabolic pool
associated with enzymatic reactions of photosynthesis, respiration, and translocation of
assimilates for tuber growth similar to that reported by Zelelew et al. (2016).
Comparing allometric functions and coefficients a and b obtained from the dry biomass
(Table 1-3) and LAI (Table 1-4) for Nc, Pc, and Kc, the present study showed that the
model based on total dry biomass was more robust in both cultivars with higher
determination coefficient for Nc, Pc, and Kc and with significant differences among the
confidence intervals (P < 0.05) of coefficients for Nc and Pc. This was because LAI was
drastically reduced at filling and maturation stage with average LAI up to 2.8 in cv. Capiro
and up to 3.8 in cv. Suprema (Figure 1-2) with loss up to 60% of leaf area starting from
flowering stage, favoring a rapid translocation of photosynthetic assimilates to the tubers,
which affects loss of turgor, specific leaf weight and, in some cases, causes leaf
abscission due to senescence, a similar phenomenon reported by Santos et al. (2010) for
cvs. Capiro and Esmeralda, and by Chakwizira et al. (2016) for crops with storage roots.
Contrary to that was reported by Zhao et al. (2014) and Ata-Ul-Karim et al. (2014) for
cereal crops, where the LAI model for Nc and Pc could be extrapolated with the function
obtained by W, probably, due to minor changes in LAI at maturity stage and minor losses
in leaf weight.
A high density of plants could increase concentrations of mineral nutrients in tissues with
higher coefficients a because it results in smaller plants (Bélanger et al., 2001; Giletto and
Echeverría 2015), but this was not the case of cvs. Capiro and Suprema (27000 plants
ha–1) with coefficients higher than 6.23-6.74 as compared to ones reported by Bélanger et
al. (2001) in cv. Shepody (44000 plants ha –1) with coefficient a around 5.36. For the
studied cultivars, a “dilution phenomenon” was more affected by the rapid growth of
tubers and translocation of assimilates to the tubers under non-limiting conditions of
fertilization and water supply than by the effects of leaf overlap, shading, or high crop
density coinciding with that reported by Justes et al. (1994) and Bélanger et al. (2001).
1.5.2 Nutrition index and optimal fertilization dose

The relationship between NI, dODF, and RY (%) followed a significant quadratic model for
cv. Capiro with most points located in deficiency or adjustment zone (NI between 0.25-0.8
and dODF < 0) and with RY between 30 and 80% in sufficiency zone (Figure 1-3), where
Nc was 2.3-2.5% between 125 to 150 dap for RY of 90-100%. The critical curves for Nc,
Pc, and Kc and NI define nutrient status of the crop, where areas of deficiency and
adjustment, sufficiency, luxury consumption or excesses could be observed as reported
by Gómez et al. (2017b).
Suprema: If NNI<1.0 than

1.2 y = -0.4862x2 + 0.802x + 0.6356
If NNI >1.0 than RY= 94%: R² = 0.36 ns
1.0
Relative Yield
0.8
0.6
0.4
Capiro:
If NNI<1.0 than y = -0.3734x2 + 1.1446x + 0.1328
0.2 If NNI>1.0 than RY = 92%
R² = 0.73***
0.0
(a) 0.00 0.50 1.00 1.50 2.00
NNI
Suprema:
1.2 y = -1E-07x2 + 6E-07x + 0.9781
R² = 0.6694**; If dOFR = 0 than RY=97.2%
1.0
0.8
Relative Yield
0.6
0.4
Capiro:
y = -4E-07x2 + 2E-05x + 1.0108
0.2 R² = 0.6547**
If dOFR = 0 than RY=95.6%
0.0
(b) -1500.0 -500.0 500.0 1500.0
dODF (kg ha-1)
Figure 1-3. Relationship between relative yield (RY) and nitrogen nutrition index (NNI) (A)
in cvs. Capiro and Suprema at tuber filling (125-150 dap) and differential optimum
fertilizer dose (dODF) (150-160 dap) (B) for cvs. Capiro ( ) and Suprema ( ) in two
production cycles in soils contrasting in fertility. ***P <0.001; *P < 0.05; ns, not significant.
In contrast, cv. Suprema presented a smaller adjustment in quadratic model with more
data located in sufficiency zone (NI 0.5-0.8), luxury consumption (NI 0.8-1.1) and areas of
excess (NI 1.2-1.5; dODF >0; RY starting from 75%), especially in soils with high N
availability (Andic Eutrudepts, Facatativá), where no response to fertilization was
observed with maximum yields lower than their production potential (Table 1-5) and to the
detriment of yield where luxury N consumption was found, limiting tuberization processes
and tuber growth. The similar data was evaluated by Gómez et al. (2017 b) for NO3- in
sap for ssp. andigena, and corroborated by Ruza et al. (2013) for ssp. tuberosum. The NI
range between 0.25 and 1.5 for the studied cultivars was lower than that reported by
Gilleto and Echeverria (2015) (NI, 0.32-1.91) and similar to those reported by Abdallah et
al. (2016) (NI, 0.39-1.53) for ssp. tuberosum.
Table 1-5. Quadratic model for yield and balanced dose of fertilizer; c: quadratic
coefficient; b: linear coefficient; a: intercept, R2: coefficient of determination. MY –
maximum yield; ODF – optimal dose of fertilizer to achieve maximum yields.
MY ODF
Cultivar Location - Cycle c b a R2 –1
(Mg ha ) (kg ha–1)
Chocontá - I -0.0000046 0.021 35.6448 0.94*** 60.6 2293

***
Chocontá - II -0.0000078 0.031 34.1819 0.86 67.7 1971
Capiro ***
Facatativa - I -0.0000300 0.059 50.9718 0.96 82.8 975
***
Facatativa - II -0.0000050 0.016 72.4079 0.98 87.0 1657
***
Chocontá - I -0.0000100 0.029 62.014 0.89 83.4 1430
***
Chocontá - II -0.0000094 0.026 47.0487 0.93 66.3 1377
Suprema
Facatativa - I -0.0000002 0.009 34.4964 ns 53.6 -
Facatativa - II 0.0000002 0.002 32.6206 ns 40.0 -
The phosphorus nutrition index and potassium nutrition index were not included because
a quadratic coefficient was positive or not significant, since the data were located mainly
in areas of sufficiency and luxury consumption, which did not allow the inflection of the
curve. It could be recommended to adjust nutrition indices of these elements with lower
doses of P and K and with less availability of these elements in soils of the Colombian
Andean region.
Regarding the optimal dose of fertilizer (ODF), a marginal response to fertilization in soil
fertility was observed for cv. Suprema without adjustment of model in two cycles with very
variable responses, where luxury consumption was proved with NI > 1 and dODF > 0.
However, in low fertility soils and at higher altitudes in Chocontá, the response was more
favorable with significance in quadratic model (Table 1-5), where ODF was established
between 1377 and 1430 kg for a maximum yield (MY) of 66.3 and 83.3 Mg ha–1 as
compared to ODF between 1971-2293 kg ha–1 in cv. Capiro that reached MY of 60.6 to
67.7 Mg ha–1 (Figure 1-3, Table 1-5).
The yield potential of cv. Suprema was more related to edaphoclimatic supply as
evidenced by the significance of location x cycle interaction, while, in cv. Capiro, it
depended more on the interaction location x fertilization, with higher expression of yield
potential in soils of high fertility (Facatativa) with ODF between 975 and 1675 kg ha –1
reaching MY of 82 to 87 Mg ha –1 (Table 1-5 and Table 1-6). It is known that fertilization
must be site-specific and cultivar-specific, such as that cv. Capiro maximum yields could
be obtained with doses 0.1-0.15 kg m–2 in high fertility soils (Facatativá) and 0.2 to 0.22
kg m–2 in low fertility soils (Chocontá), while cv. Suprema in soils of lower fertility requires
0.135-0.145 kg m–2, that is 30% less fertilization than cv. Capiro, these ranges were lower
than those reported by Gómez and Torres (2012) (0.2 and 0.3 kg fertilizer m –2).
Table 1-6. Average yield of tubers (Mg ha –1) harvested at 150-160 dap for cvs. Capiro
and Suprema and statistical significance of mean square of factors and their interactions.
Factor Tuber fresh weight (Mg ha–1)
Cycle Capiro (A) Suprema (B)
I 60.56 (b) 69.31 (a)
II 78.76 (a) 47.55 (b)
Location
Chocontá 59.17 (b) 67.26 (a)
Facatativá 80.16 (a) 49.60 (b)
Fertilization
F0 52.08 (b) 65.25 (a)
F1 78.77 (a) 59.35 (a)
F2 75.98 (a) 56.65 (a)
F3 71.82 (a) 52.47 (a)
Interactions Mean squares
Cycle 2391.1*** 4392.8**
Location 7330.9*** 4934.4***
Fertilization 1993.2*** 331.5 ns
Cycle x Location 1664.7*** 1135**
Cycle x Fertilization 144.9 ns 156.9 ns
Location x Fertilization 321.4*** 241.9 ns
Cycle x Location x Fertilization 144.4 ns 31.27 ns
Error 9119.0 9120
Similar letters indicate non-significant statistical differences per cultivar (lowercase) and between
cultivar (capital letters) according to the LSD test, P < 0.05. **Significant P < 0.05; ***highly
significant P<0.001.
1.6 Conclusions
The allometric relationships by dilution curves for Nc, Pc, and Kc were quantified for the
first time in cvs. Capiro and Suprema of Group Andigena. The Nc, Pc, and Kc of best
adjustment for cvs. Capiro and Suprema were those obtained from total dry biomass W
with respect to LAI due to the higher variation in leaf area, a product of rapid translocation
of photoassimilates to the tubers. The studied cultivars only had differences in critical
dilution curve for Kc, where cv. Capiro presented a higher requirement Kc at initial stages
of growth than cv. Suprema demonstrating, with higher coefficient a and lower coefficient
b, that cv. Capiro was more efficient in use of K to obtain higher dry biomass. For the first
time, critical dilution curves for Nc, Pc, and Kc were obtained for potato Group Andigena
making it one of the first references of Kc for potato. The critical dilution curves were
adjusted to the model proposed for Group Chilotanum but with less dilution in coefficients
a and b for N and P, which shows a higher requirement of these elements with lower
nutrient efficiency. The highest adjustment of NI in the studied areas showed that N was
the most limiting element for obtaining high yields, mainly in cv. Capiro. In addition, cv.
Suprema was more prone to luxury uptake and doses in soils of low fertility can be 70% of
the optimal dose for cv. Capiro. Determining nutritional status of N, P, and K at early
growth stages from dilution curves allows providing timely adjustments in management of
these elements at tuberization and filling stages and, thus, potentiates the yields.
Acknowledgements
The authors express their gratitude to INGEPLANT SAS, FEDEPAPA, and Universidad
Nacional de Colombia-Bogotá for the funding and support in laboratory analysis. The
authors acknowledge the help of research assistants Mrs. Paola Torres, Mrs. Liliana
Arevalo, Mr. Elías Silva, and Mrs. Andrea Barragán for field support. We also thank the
growers Mr. Walter Guzmán – Biogenética, Mr. Yovanny Pulido, and Mr. Ricardo Rojas
for the logistical contribution to the development of this research.
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2. Uptake and partition of N, P and K in potato
(Solanum tuberosum L. Group. Andigena)1
Consumo y partición de N, P y K en el cultivo de papa

(Solanum tuberosum L. Grupo Andigena)
Manuel Iván Gómez, Stanislav Magnitskiy, Luis Ernesto Rodríguez
2.1 Abstract
Uptake curves for nitrogen (Nuptake), phosphorus (Puptake), and potassium (Kuptake) obtained
from total dry biomass (W) together with analysis of nutrient allocation into the tubers
derived from nutrient harvest index (NHI, PHI, KHI) are important for prognosis of potato
growth in the Andean region. These indices permit a more specific management of
mineral nutrients according to soil type and plant genotype obtaining maximum yields at
adequate fertilizer doses and avoiding economic and environmental impacts on potato
production systems. This research established critical uptake curves (N uptake, Puptake, and
Kuptake) and translocation curves for N, P, and K using allometric relationships in potato
cultivars Diacol Capiro and Pastusa Suprema (Solanum tuberosum, Group Andigena)
cultivated on soils of high and low fertility. The significant differences were observed in the
genotype x location interaction with the best fit of uptake model obtained on low fertility
soils for Suprema (Nuptake = 68.13W0.504, Puptake = 6.724W0.779, and Kuptake = 63.93W0.776),
where this cultivar expressed the highest productive potential and high NHI (0.55-0.69),
PHI (0.75-0.8) and KHI (0.62). Capiro was better adapted to the changes in soil fertility
than Suprema, since the models for nutrient uptake in Capiro were significant for both
soils. The efficiency of N, P, and K translocation into the tubers followed a positive
logarithmic model and revealed the differences in favor of Capiro in efficiency of
1
Sometido en Field Crops Research Journal
partitioning of assimilates and N, P, and K. The uptake of mineral nutrients was identified
per cultivar and per phenological stage, with up to 33% more P extracted by tubers in
Capiro than in Suprema and comparable extraction of N and K by tubers in both cultivars.
The uptake of these elements was differentiated by phenological stage permitting a more
precise adjustment of fertilization plans.
Key words: harvest index, efficiency of translocation, uptake curves, macronutrient
extraction
2.2 Resumen
Las curvas críticas de consumo de nitrógeno (Nr), fósforo (Pr) y potasio (Kr)
determinadas a partir de la biomasa seca total (W) y el análisis de la partición nutricional
en el tubérculo y el índice de cosecha de nutrientes (ICN, ICP, ICK) son importantes para
cultivares de importancia económica del país, porque permite evaluar las diferencias
entre genotipos y se puede realizar un manejo más específico de nutrientes por tipo de
suelo, cultivar y fenología en busca de disminuir la brecha tecnológica para alcanzar
máximos rendimientos con un uso adecuado de los fertilizantes y así evitar impactos
económicos y ambientales en sistemas productivos de papa. El objetivo de este estudio
fue determinar curvas de consumo de Nr, Pr y Kr (kg ha-1) y curvas de traslocación de
nutrientes (ENt, EPt y EKt) mediante relaciones alométricas durante el crecimiento en
dos ciclos de cultivo en cvs. Diacol Capiro (Capiro) y Pastusa Suprema (Suprema) bajo
suelos alta y baja fertilidad. Se observó una significancia en la interacción genotipo x
localidad con el mejor ajuste del modelo de consumo en suelos de baja fertilidad para
Suprema (Nr = 68,13 W0,504, Pr = 6,72 W0,779 y Kr = 63,93 W0,776) donde expresa el mayor
potencial productivo y altos ICN (0,55-0,69), ICP (0,75-0,8) e ICK (0,62), mientras Capiro
muestra una mayor adaptación en ambos tipos de suelos porque los modelos de
consumo fueron significativos con mejor significancia para Nr = 56,38 W0,58 y para Pr =
4,26 W0,786 en suelos de menor fertilidad y para Kr = 79,52 W0,79 en suelos fértiles. La
eficiencia de traslocación sigue un modelo logarítmico positivo y evidencia diferencias a
favor de Capiro en la partición de asimilados y nutrientes con mejor eficiencia en el uso
de N, P y K por su hábito determinado. Se estableció requerimientos nutricionales por
cultivar y por etapa fenológica con un 35% más de extracción nutricional de P en Capiro
que en Suprema y similares de N y K, con diferenciación en consumo por fenología lo
cual permite realizar ajuste más precisos en planes de fertilización.
2. Uptake and partition of N, P and K in potato (Solanum tuberosum L. Group. 41
Andigena)
Palabras clave: índice de cosecha de N, P y K, eficiencia de translocación, curvas de

consumo, extracción nutricional.
2.3 Introducción
The distribution of assimilates among sink-source organs determines crop yield (Gifford
and Evans, 1981) and, in potato (Solanum tuberosum L.) production, is strongly regulated
by water and nutrient availability (Schilling et al., 2016). The relationships between
accumulation of nutrients in different organs with respect to increment of plant dry mass
(Zhao et al., 2017) reflect requirements of mineral elements that plants need for adequate
growth and development. Although uptake of mineral nutrients with respect to
accumulation of biomass in potato is genetically determined (Bélanger et al., 2001;
Sandaña and Kalazich, 2015), it could be modified by environmental factors (Ata-Ul-Karim
et al., 2014), fertilizer dose (Swain et al., 2014), and initial soil fertility (Gómez et al.,
2017).
The critical curve of nutrient uptake Nuptake is important to predict the amount of a mineral
nutrient needed to produce a given biomass (Andriolo et al., 2006) according to crop
production potential and is defined by the following allometric relationship adapted from
Bélanger et al. (2001) and Cogo et al. (2006):
Nuptake (kg ha-1)=aWb (equation 1),
where W is total biomass (Mg ha-1), Nuptake is uptake of N (Nuptake), P (Puptake), or K (Kuptake)
(kg ha-1) by plants at a given total biomass. The coefficient a represents the concentration
of nutrient when total biomass is equal to or exceeds 1 Mg ha -1, while parameter b
represents the "uptake coefficient" that determines the differences in nutrient
requirements as long as the total biomass is increasing.
According to Lemoine et al. (2013) and Griffith et al. (2016), assimilate partitioning in
green plants starts with: (i) photosynthetic activity (carbon fixation) in leaves, (ii)
conversion of carbon fixed in leaves into sucrose and synthesis of starch in chloroplasts,
(iii) sucrose loading into the phloem, (iv) long-distance translocation of sucrose, and (v)
phloem unloading and use of sucrose by source organs. Assimilation of carbon in plants
is coupled to assimilation of nitrogen to ensure that basic components of biomass
production, such as amino acids and carbon skeletons, are available in the required
quantities and stoichiometry, and to their transport from the sites of synthesis to the sites
of demand. Carbon gain by plants depends intrinsically on the successful absorption of

other mineral nutrients, particularly, N and P (Griffith et al., 2016) and K (Lemoine et al.,
2013).
Compartmentalization of mineral nutrients in leaves, stems, and tubers during potato

growth could be used to define the nutrient status of the crop. This process describes
nutrient allocation within the plant and reflects the differences existing between the
cultivars (Swain et al., 2014) by means of nutrient harvest index (Giletto and Echeverría,
2015) and nutrient translocation efficiency (Fernandes and Soratto, 2012) as indicators of
partitioning of assimilates and mineral nutrients. In potato, these indexes describe well the
nutrient uptake under conditions of nutrient or water deficit (Schilling et al., 2016),
contrasting temperatures, and changes in photoperiod (Wolf et al., 1990) or soil fertility
(Gómez et al., 2017). Additionally, the stage of plant development constitutes a factor that
affects partitioning of mineral elements, with a high export of assimilated elements from
potato leaves after formation of tubers (Wolf et al., 1990).
In the world, potato is one of the major crops defining food security and reducing poverty
and human malnutrition (George et al., 2018). At the same time, in potato Group
Andigena, which represents the second largest product of the basic consumer basket in
Colombia, almost no studies were done in uptake of mineral elements by the plants
(Gómez et al., 2017). The objective of this study was to evaluate uptake and partitioning
of primary macronutrients N, P, and K in two potato cultivars Diacol Capiro (Capiro) and
Pastusa Suprema (Suprema) (Andean cultivated tetraploids Solanum tuberosum L.,
Group Andigena) at different phenological stages of growth. The research aimed to
establish the uptake curves for N (Nuptake), P (Puptake), and K (Kuptake) and their translocation
efficiency under fertilization on soils contrasting in fertility. The study planned to evaluate
partitioning of N, P, and K by means of harvest indexes that would allow to characterize
the cultivars and, thus, to predict nutrient requirements and time-specific management of
these macronutrients in field.
2.4.1 Experimental locations and conditions

The research was carried out in two production cycles (2013-2016) in two locations of the
Andean region of Colombia: (i) Facatativá with Andic Eutrudepts soils, highly fertile and
Andigena)
saturated in bases, and (ii) Chocontá with Humic Dystrudepts soils of low fertility, acid and
desaturated in bases. These locations were representative for the region of potato
production in Colombia and had contrasting edaphoclimatic conditions (Table 2-1).
Table 2-1. Characteristics of climate and soils in experimental locations.

Annual Rainfall T T
Altitude North West Evaporation T min
1
Climatic Crop rainfall (mm -1
max media
(m.a.s.l.) latitude longitude -1
(mm cycle ) (ºC)
variables cycle (mm) cycle ) (ºC) (ºC)
Cycle 1 4° 49´ 74° 22’

2,520 951 397 454 18.1 7.0 12.7
(I-2013) 26.9” 29.7”
Facatativá
Cycle 2
4° 49´ 74° 22’
(II- 2,520 850 415 382 18.5 7.2 12.5
26.9” 29.7”
2015)
Cycle 1
5° 5´ 73° 43´
(II- 2,780 1,295 712 640 16.2 4.4 10.6
30.37” 2.04”
2013)
Chocontá
Cycle 2 5° 5´ 73° 43´
2,710 1,058 803 603 16.5 10.1 12.98
(I-2016) 30.37” 2.04”
CEC K
Soil Al (cmolc O.M. (g -1
P (mg (Ca +
Texture pH -1 -1
(cmolc N (g kg ) -1
(cmolc
2 fertility †† kg ) kg ) kg ) Mg)/K
Soil kg )
-1
kg )
-1
characteristics
Facatativá Loam High 6.4 0 166.7 31.95 8.3 39.64 3.14 9.1
(Andic
Eutrudepts) Loam High 5.82 <0.001 127.1 19.14 6.4 70.16 0.87 20.87
Clay
Chocontá Low 5.5 0.1 67.7 9.52 3.3 18.18 0.68 12.8
Loam
(Humic
Clay
Dystrudepts) Low 5.28 0.54 85.9 7.9 4.3 41.5 0.84 7.56
Loam
1
Climatic data obtained and calculated from IDEAM, Colombia (IDEAM, 2017). 2 Physical-chemical
characteristics of soils and potential chemical fertility determined in arable layer (0-30 cm) (IGAC,
2006; Castro and Gómez, 2013). The soils were classified according to USDA (Soil Survey Staff,
2010).
An incomplete factorial arrangement was used in divided subplots of a mixed model with
four replicates (one plant per replicate) arranged in randomized complete block design
with main plot corresponding to the cultivars (Capiro and Suprema) and the subplots
matching to four levels of fertilization (F0, F1, F2, and F3); F0 referred to unfertilized plots
(initial conditions of soil fertility). Intra-subject factor of time was associated with five
critical phenological stages of crop growth adapted from Valbuena et al. (2010): Stage I,
50-55 days after planting (dap) (formation of primary stems and ramification); Stage II, 70-
75 dap (formation of secondary stems and initiation of tuberization); Stage III, 90-100 dap
(flowering, maximum tuberization, and start of tuber filling); Stage IV, 120-125 dap
(ceasing of flowering and tuber filling); Stage V, 150-160 dap (senescence, maximum
tuber filling, and tuber maturation). The contribution of mineral nutrients through fertilizer
plans, fertilizer sources, and dose fractionation are presented in Table 2-2.
Table 2-2. Contribution of mineral nutrients through fertilizer applications.
Crop 1
Fertilizer dose Supply of mineral nutrients (kg ha-1)
Location -1
cycle (kg ha )
N P2O5 K2O Mg S B Zn Mn
F1 1,186 128 196 135 53 56 2.55 4.2 5.25
Cycle 1 F2 1,582 171 261 180 70 74 3.4 5.6 7.0
F3 1,977 214 326 225 88 93 4.3 7.0 8.8
Facatativá
F1 1,450 123 216 176 60 113 1.7 3.5 4.2
Cycle 2 F2 1,900 164 288 235 80 150 2.3 4.6 5.6
F3 2,375 205 360 294 100 188 2.9 5.8 7
F1 1,632 144 255 261 42 90 0.9 1.8 2.3
Cycle 1 F2 2,175 192 340 348 56 120 1.2 2.4 3.0
F3 2,719 240 425 435 70 150 1.5 3.0 3.8
Chocontá
F1 1,500 143 285 236 30 29 3.3 3.6 4.1
Cycle 2 F2 2,000 191 380 315 40 38 4.4 4.8 5.4
F3 2,500 239 475 394 50 48 5.5 6.0 6.8
1
Fertilizer doses suggested using a soil-plant balance method (Castro and Gómez, 2013); fertilizer
doses fractionated following the historical practices employed in locations of the highest yields
-1
(>50 Mg ha ): N 60% at planting, 40% 45-50 dap; P 70% at planting, 30% 45-50 dap; K 30% at
planting, 70% 45-50 dap. Granulated fertilizers were used: DAP for N and P; KCl (0-0-60) and
K2SO4 (0-0-50) for K; Ca(NO3)2 (25% CaO) for Ca; MgSO4·H2O for Mg; a complex micronutrient
fertilizer based on sulfates Nutricomplet® (Ingeplant, Colombia) for B, Zn, Cu, Mn, and Fe.
The tubers of about 70 g were planted in experimental units of 50 m 2 (135 plants plot-1),
with a distance 1 m between the rows and 0.37 m between the plants, a useful area of 36
m2 (three central rows) and a density of 27,000 plants ha -1. The practices of irrigation,
weed management, and phytosanitary control were scheduled when required. For
Chocontá location of highly acidic soils, a dolomite-type amendment was applied preplant
at a dose of 1.0 Mg ha-1.
Andigena)
2.5 Analytical methods

In each production cycle, four plants were selected per experimental unit and per growth
stage and a destructive analysis of leaves, aerial stems, and tubers was performed. For
chemical analysis, plant organs were rinsed with deionized water, then each organ was
mixed up to obtain a subsample of 200 g (fresh weight) and dried in an oven at 70 °C for
72 h, after that the dry weight (DW) was measured. The total concentrations of N, P, and
K were determined in each organ according to IGAC (2006). The total contents of N, P,
and K in each organ were calculated by multiplying a concentration of mineral nutrient in
the given organ (g 100 g-1 DW) by its dry biomass W (kg ha-1) at a respective
phenological stage (Abdallah et al., 2016). The differences among distribution of
assimilates within potato plants were evaluated by indices of physiological efficiency of
mineral nutrients in potato adapted from Fernandes and Soratto (2012) and Giletto and
Echeverría (2015):
Efficiency of translocation of N (EtN), P (EtP), or K (EtK) into the tubers was evaluated by
determining nutrient accumulation in tubers Nut (kg ha-1) with respect to total nutrient
uptake Nuptake (kg ha-1) at given moment of growth: EtN = (Nut / Nuptake) x 100%.
Harvest index HI was calculated as a relation between dry biomass of tubers DWt and
total (aerial part + tubers) dry biomass W of the plant: HI = (DWt / W).
Nutrient harvest index for N (NHI), P (PHI), or K (KHI) was calculated as a relation
between nutrient uptake by tubers (Nut, kg ha-1) at 150-160 dap and total (aerial part +
tubers) nutrient uptake by the plant (Nuptake, kg ha-1): NHI = (Nut / Nuptake). 
2.5.1 Statistical analysis

A multivariate analysis was carried out for cultivar, location, fertilizer dose, and crop cycle
at each sampling moment (dap) with respect to total dry biomass W (kg ha-1). When
significant differences were found between the fertilizer doses in individual analysis (Table
2-2), LSD test (p <0.05) was utilized to select the data with nutrient uptake corresponding
to maximum W values. From the selected data, a non-linear regression was obtained to
adjust the data to the allometric curve. The uptake curves for N uptake, Puptake, and Kuptake (kg
ha-1) with respect to the total biomass W were proposed according to equation 1 using the
potential models of Bélanger et al. (2001) and Cogo et al. (2006). The curves of nutrient
translocation efficiency EtN, EtP, and EtK were determined over time using a logarithmic
regression analysis. The coefficient of determination (R 2) was calculated from the relation
between the sum of squares of the error and the total sum (adjusted coefficient), and the
95% confidence intervals were constructed for each of the estimators (a and b) to
evaluate differences of models between the cultivars or the most significant factors.
For biomass and nutrient harvest indexes (NHI, PHI, and KHI) at 150-160 dap, a mixed
model was used assuming the replicates to be random effects, while the main effects,
cultivar, location, crop cycle, and fertilizer levels were assumed to be fixed effects.
Considering all interactions between the main effects, the different interactions between
the factors were evaluated by LSD test (p < 0.05); the mean comparison (LSD) was
performed for a statistically significant interaction of the highest level in this case. For all
statistical procedures described previously, SAS 9.4 software (SAS Institute, 2014) was
used. Because the interaction genotype x environment was determined to be highly
significant, analysis of covariables of soil fertility and climate was performed to determine
which one(s) accounted for a significant proportion of the variability of the interaction, for
variables of nutrient uptake, total yield, and total dry biomass. This analysis was
performed by GEAR software (Pacheco et al., 2015).
2.6 Results
2.6.1 Total uptake of N, P, and K and nutrient harvest indexes

According to the ANOVA, a highly significant interaction location x cultivar (p < 0.001) was
determined for the means of nutrient uptake (N uptake, Puptake, and Kuptake) at different
phenological stages; no effects were revealed for the factors “production cycles” and
“levels of fertilization”, which explains that the total uptake of N, P and K in Capiro and
Suprema was mostly mediated by a genotype-environment factor.
Dynamics of total biomass accumulation (W, Mg ha−1) per phenological stage and
potential tuber yield (PYt, Mg ha −1) were affected mostly by soil fertility (p <0.001) than by
climatic factors according to analysis of covariance carried out in two locations and
production cycles. For this, a model searched to describe a behavior of total nutrient
uptake with respect to total biomass accumulation by comparing soil fertility in each
studied location (Figure 2-1). Figure 2-1 shows close allometric relations among total dry
biomass (W) and uptakes of N, P, or K during crop growth using a logarithmic model,
Andigena)
which accords to the models for N uptake in potato (Bélanger et al., 2001) and rice (Oryza
sativa L.) (Ata-Ul-Karim., 2014) and for K uptake in potato cv. Asterix (Cogo et al., 2006).
1000
1000 Low fertility High fertility Nuptake = 78.018W0.6918
(a) 900 (b)
R² = 0.67786*
N uptake cv. Suprema (kg ha-1)

N uptake cv. Capiro (kg ha-1)
Nuptake = 73.594W0.6588 800

800 R² = 0.8246**
700
600
600
500
400 400
300
200 200
Nuptake = 56.383W0.584 Nuptake = 68.131W0.5048
R² = 0.9033** 100
R² = 0.8682**
0 0
0 5 10 15 20 25 30 35 0 5 10 15 20 25 30 35
100 (c) 100
P uptake cv. Suprema (kg ha-1)

90
P upttake cv. Capiro (kg ha-1)
90 Puptake = 6.5042W0.7538 Puptake = 6.7246W0.7795

80 R² = 0.9181** 80 R² = 0.89419** (d)
70 70
60 60
50 50
40 40
30 30
20 Puptake = 4.2675W0.786 20
R² = 0.96908** Puptake = 4.7655W0.6625
10 10
R² = 0.87755**
0 0
0 5 10 15 20 25 30 35 0 5 10 15 20 25 30 35
1400 1400
K uptake cv. Capiro (kg ha-1)
Kuptake = 79.521W0.799
K uptake cv. Suprema (kg ha-1)
1200 (e) 1200 (f)

R² = 0.85355** Kuptake = 63.934W0.7761
1000 1000 R² = 0.91032**
800 800
600 600
400 400 Kuptake = 42.774W0.7724
200 Kuptake =
77.637W0.6713 200 R² = 0.39325 ns
R² = 0.7595*
0 0
0 5 10 15 20 25 30 35 0 5 10 15 20 25 30 35
Total dry biomass (Mg ha-1) Total dry biomass (Mg ha-1)
Figure 2-1. Average total uptake of N, P, and K in cvs. Capiro (left) and Suprema (right)
as a function of total dry biomass (W) in soils of high (Andic Eutrudepts, Facatativa) and
low (Humic Dystrudepts, Chocontá) fertility and two production cycles under non-limiting
conditions of fertilization. **Indicates significant values of coefficients b with respect to soil
fertility at a probability level of 0.05. ns, non-significant model.
The prominent slopes in uptake curves Nuptake and Puptake for both cultivars indicated the
presence of significant differences (p<0.05) among the coefficients b for soil fertility
(Figure 2-1, Table 2-3), where the demand of nutrients in relation to total biomass
depended mainly on the high fertility of Andic Eutrudepts as compared to the low fertility
of Humic Dystrudepts (Table 2-1). However, the values of Kuptake in Suprema on fertile
soils did not result in significant differences (R 2 = 0.39), while the model for K uptake in
Capiro on fertile soils was highly significant (Figure 2-1). Additionally, for Nuptake in
Suprema, a significance (R2 = 0.68) was lower than the one found in Capiro (R 2 = 0.82).
120
110 Capiro Suprema
100
Potential yield (Mg ha-1)
90
80
70
60
50
40
30
20
10
0
Chocontá - 1 Facatativa - 1 Chocontá - 2 Facatativa - 2
Location - Cycle
Figure 2-2. Potential yield (PYt) in cvs. Capiro and Suprema cultivated in soils of low
(Humic Dystrudepts, Chocontá) or high (Andic Eutrudepts, Facatativá) fertility at 150-160
dap under non-limiting conditions of fertilization in two production cycles. P<0.05 for
fertilization x location x cultivar. Error bars indicate standard errors.
1 Capiro Suprema
0.9 a a
b b b b
0.8
0.7
Harvest index
0.6
c
0.5
d
0.4
0.3
0.2
0.1
0
Chocontá 1 Facatativa 1 Chocontá 2 Facatativa 2
Location x cycle
Figure 2-3. Harvest index (HI) in cvs. Capiro and Suprema cultivated in soils of low
(Humic Dystrudepts, Chocontá) or high (Andic Eutrudepts, Facatativá) fertility under non-
limiting conditions of fertilization in two production cycles. P<0.05 for location x cultivar x
cycle. Means followed by the different letters are significantly different (LSD, P<0.05).
Error bars indicate standard errors.
Andigena)
Suprema had better adaptation to desaturated acid soils of higher altitudes under non-
limiting conditions of fertilization agreeing with its genotypic features of longer stolons and
a deeper root system that improves absorption of mineral nutrients as shown by Valbuena
et al. (2010). Low fertility soils for Suprema favored tuberization and ensured a better
yield of 63 and 73 Mg ha −1 when compared to 46-50% losses in productive potential on
high fertility soils and yields of 31-41 Mg ha−1 (Figure 2-2). Furthermore, high soil fertility
favored a continuous allocation of N and K into aerial parts of Suprema plants similar to
that observed in Capiro during tuber filling and maturation. However, this was to the
detriment of biomass accumulation in tubers with reduced translocation efficiency of
assimilates evidenced by low HI (40 to 50%) for Suprema as compared to HI 75-78% for
Capiro in the same soil (Figure 2-3).
Suprema presented a higher requirement in N when W was equal to or less than 1 Mg

ha−1 (early growth stage, 55-60 dap), with 78.01 and 68.13 kg N ha -1 uptake on high and
low fertility soils, respectively. In this case, for Suprema, the values of coefficient a were
higher and significantly different than the ones in Capiro, which had 73.59 and 56.38 kg N
ha-1 uptake on high and low fertility soils, respectively. This could be due to a higher N
requirement for superior biomass of branches and stems in Suprema as compared to
Capiro. This advocates for a timely supply of N to the plants before tuberization (45 dap)
at a rate of 10-15% of its total uptake in Suprema and 20-24% in Capiro.
Significant differences among the cultivars were found in "uptake coefficient" b for Nuptake
on high fertility soils (p <0.05) indicating a higher demand for N in Suprema due to a
higher W with b = 0.69 with respect to b = 0.65 in Capiro (Figure 2-1). However, Suprema
had low productive potential (Figure 2-2) with HI ranging from 0.4 to 0.5 (Figure 2-4A) that
were lower when compared to HI of 0.75-0.8 in various accessions of Group Andigena
(Zebarth et al., 2008). At the same time, in poor soils of Chocontá, no significant
differences were found between b of 0.58 and 0.5 for Capiro and Suprema, respectively,
with similar N uptake in these cultivars at the highest W. This high uptake of N was
related to a better yield potential (Figure 2-2) and adequate HI of 0.75 to 0.82 in both
cultivars (Figure 2-3), which illustrates a highly productive potential of potato Group
Andigena as reported by Zebarth et al. (2008).
0.9 Capiro Suprema (a)

a
0.7 ab
b ab
b b
N Harvest Indiex
0.5
c
0.3 c
0.1
-0.1 Chocontá 1 Facatativa 1 Chocontá 2 Facatativa 2
0.9 a a a (b)
a a a
0.8
0.7
P Harvest Index
0.6
0.5 b
0.4 b
0.3
0.2
0.1
0
0.9 (c)
0.8 a a
a a
0.7
b
K Harvest Index
0.6 bc
c
0.5
0.4
0.3 d
0.2
0.1
0
Location x cycle
Figure 2-4. Harvest index for N (NHI) (A), P (PHI) (B), and K (KHI) (C) in cvs. Capiro and
Suprema cultivated on Humic Dystrudepts (Chocontá) and Andic Eutrudepts (Facatativá).
P<0.001 for location x cultivar x cycle under non-limiting conditions of fertilization. Means
followed by the different letters are significantly different (LSD, P<0.05). Error bars
indicate standard errors.
Andigena)
A difference in uptake curves was obtained between the cultivars for P (P uptake) (Figure
2-1C, D), with a equal to 6.5 and 4.7 kg P ha -1 and b equal to 0.75 and 0.66 in Capiro and
Suprema, respectively, on fertile soils. In contrast, Suprema presented larger P uptake at
initial stages of growth with a = 6.72, while Capiro had a = 4.26 kg P ha-1 and no
significant difference (p > 0.05) in b were found among genotypes in poor soils. This
indicated similar P requirements in both cultivars, with efficient translocation of P to tubers
and PHI ranging between 0.75 and 0.8 (Figure 2-4B). The high accumulation of P in
tubers was due to its role of an essential element in formation of amylopectin and
amylase for the synthesis of starch, which represents 15 to 20% of the tuber weight
(Chung et al., 2014; Leonel et al., 2016).
In both locations, the coefficient a for Kuptake presented significant differences favoring
Capiro and reaching 79.52 and 77.63 kg ha-1 on fertile and poor soils, respectively,
whereas, in Suprema, a was equal to 42.7 and 63.9 kg ha-1 on fertile and poor soils,
respectively (Figure 2-1E, F), which revealed a higher K requirement in Capiro at initial
stages of growth. This finding was, probably, due to earlier tuberization and tuber filling in
this cultivar. In contrast, on poor soils, Suprema presented differences (p <0.05) in
coefficient b = 0.77 that surpassed the one in Capiro with b = 0.67. Therefore, in order to
reach W of 25 Mg ha−1 (PYt of 80 Mg ha−1), a total K uptake of 762 kg ha -1 and of 673 kg
ha-1 was needed in Suprema and Capiro, respectively. Capiro was more efficient in the
use of K producing tubers with a less variable KHI (0.5-0.7) than Suprema (0.13-0.62) on
both low and high fertility soils (Figure 2-4C).
Considering uptake curves (Figure 2-1) and HI (Figure 2-4) on soils of highest productive
potential, the total uptake and extraction by tubers of N, P, and K could be predicted for a
given level of production (Table 2-3). This would allow adjustment of the fertilizer
recommendations for the projected yields in Capiro and Suprema. Additionally, 2.01 and
1.76 kg of K were absorbed by Suprema and Capiro, respectively, per 1 kg of N as
compared to 1.9 kg K absorbed per 1 kg of N in cv. Asterix (Group Chilotanum) (Cogo et
al., 2006). A 10-12% higher extraction for N and K and 33% higher extraction for P were
obtained in Capiro compared to Suprema in order to reach high (> 65 Mg ha −1) yields
(Table 2-3).
Table 2-3. Prognosis of total uptake and extraction by tubers for N, P and K in cvs. Capiro
and Suprema at different yields in soils of high yield potential of the Colombian Andean
region.
2
1
Extraction by Extraction index EI
Potential yield Total uptake
W tubers (kg of nutrient
Cultivar (PYt) (kg ha-1) -1
(kg ha ) Mg-1 PYt)
-1 -1
Mg ha Mg ha N P K N P K N P K
15 5.0 154 14 223 83 10 123 5.6 0.70 8.2
32 10.0 218 22 382 118 16 210 3.7 0.51 6.5
Suprema 49 15.0 267 29 523 144 21 288 2.9 0.44 5.8
66 20.0 309 35 654 167 26 360 2.5 0.39 5.4
84 25.0 346 40 777 187 30 428 2.2 0.36 5.1
17 5.0 144 15 229 82 11 160 4.9 0.68 9.6
35 10.0 216 26 364 123 20 255 3.5 0.56 7.3
Capiro 54 15.0 274 36 478 156 27 335 2.9 0.50 6.3
72 20.0 324 45 580 185 34 406 2.6 0.47 5.6
90 25.0 369 54 674 211 40 472 2.3 0.44 5.2
3
Percentage of
increment, Capiro 7.96 0.0 6.79 33.26 -13.34 12.72 33.26 10.30 4.41 23.44 2.17
vs Suprema
1
The quantities (kg ha-1) estimated for Capiro: N = 56.38W0.584; P = 4.27W0.786; K = 77.63W0.786,
and for Suprema; N = 68.131W0.505; P = 6.724W0.779; K = 63.934W0.786. The yield (Mg ha-1) was
estimated from total dry biomass (W) in Capiro (W = PYt 0.2717 + 1.6959) and Suprema (W = PYt
0.2908 + 2.311). 2 Nutrient extraction by tubers estimated from NHI, PHI, KHI in each cultivar
(Figure 2-4). 3 The comparison done between total uptake and extraction by tubers of mineral
elements in Suprema with respect to Capiro (% of relative increase) at a projected W of 25 Mg ha-
1
.
2.6.2 Efficiency of translocation and extraction of N, P, and K by

tubers
The timely changes in the efficiency of nutrient translocation in plants followed a positive
logarithmic model (Figure 2-5) with highly significant differences (P <0.001) observed in
the interaction genotype x location, which indicated that partitioning of assimilates in
potato was affected by the stage of tuber development as referenced by Wolf et al.
(1990).
Andigena)
100 Suprema EtN = 54.498ln(dds) - 220.62 ns, R² = 0.991 A
Efficiency of N traslocation (%)

90
Capiro EtN= 55.866ln(dds) - 228.82 ns, R² = 0.964
80
70
60
50
40
30
20
10
0
0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160
100 Suprema EtP = 60.833ln(dds) - 244.36**, R² = 0.998 B

Efficiency of P traslocation (%)
90
80 Capiro EtP = 77.898ln(dds) - 315.47**, R² = 0.9895
70
60
50
40
30
20
10
0
0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160
100 C
Suprema EtK = 49.035ln(dds) - 197.64**, R² = 0.993
Efficiency of K traslocation (%)
90
80 Capiro EtK= 62.929ln(dds) - 255.62**, R² = 0.981
70
60
50
40
30
20
10
0
0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160
Days after planting (dap)
Figure 2-5. Average translocation efficiency of N (EtN), P (EtP), and K (EtK) in cv. Capiro
(circle) and cv. Suprema (square) in two production cycles under non-limiting conditions
of fertilization in contrasting soils. ** Significant differences of coefficients b are shown
with respect to the interaction cultivar x location at a probability level of P <0.05.
On the other hand, a lower EtN (7-10%) and reduced extraction of N were observed at the
start of tuberization indicating a lower N:P:K ratio in Suprema than in Capiro (Table 2-4),
with lesser uptake of N favoring tuber differentiation in Suprema. These results were
similar to that reported by Fontes et al. (2016) for cvs. Asterix and Atlantic.
Table 2-4. Extraction of N, P, and K by tubers per growth stage for cvs. Suprema and
Capiro. The values are estimates from translocation efficiency of N (EtN), P (EtP), and K
(EtK) with a projected yield of 70 Mg ha −1.
Extraction by Accumulation per

-1
tubers (kg ha ) stage (% of the
Phenological stage Ratio N:P:K
Cultivar for yield of 70 extraction by
-1
Mg ha tubers)
Stage dap N P K N P K N P K
Start of
55-75 25 7 75 15 26 20 7 2 22
tuberization
Flowering,
maximum 75-100 82 16 217 48 58 58 11 2 28
tuberization
Tuber filling,
Suprema
cessation of 100-125 123 22 276 72 82 74 11 2 25
flowering
Maximum
filling,
125-160 171 27 373 100 100 100 13 2 28
maturation of
tubers
Start of
55-75 38 8 78 21 22 20 10 2 20
tuberization
Flowering,
maximum 75-100 93 20 216 51 55 54 9 2 22
tuberization
Tuber filling,
Capiro
cessation of 100-125 140 29 331 77 82 83 10 2 23
flowering
Maximum
filling,
125-160 182 36 399 100 100 100 10 2 22
maturation of
tubers
Andigena)
2.7 Discussion
2.7.1 Total uptake of N, P, and K and nutrient harvest indexes

The results obtained for nutrient uptake (Figure 2-1) were similar to that described for N
by Bélanger et al. (2001) in cvs. Shepody and Russet Burbank (Group Chilotanum) and
contrary to that identified by Swain et al. (2014), when the interaction genotype x
fertilization (conventional vs. organic) affected uptake and efficiency of N use in cvs.
Sarpo Mira and Sante (Group Chilotanum). On the other hand, Fernandes et al. (2017)
found that increasing doses of P improved uptake of this element by potato plants, with
differences obtained between the genotypes of Group Chilotanum that favored cv.
Mondial with respect to cv. Agata.
Capiro was more efficient in allocation of assimilates to the source organs on fertile soils,
with yields ranging from 70 (Cycle I) to 99 (Cycle II) Mg ha −1 (Figure 2-2) and high HI of
78 and 83% (Figure 2-3). This might be due to a compact size of Capiro plants of
determined growth habit, shorter stolons, and limited period of tuberization between 75-
100 dap, the conditions that favored better translocation of assimilates and faster growth
after tuber formation as reported in Chilotanum cultivars by Gliletto and Echeverría
(2015). The differences in yield and dry biomass and high HI for Capiro were determined
mainly by a higher soil offer of P (p < 0.001) and an improved ratio Ca+Mg/K (p < 0.001)
with better availability of soil K (Table 2-1) according to analysis of covariance.
Additionally, Fernandes et al. (2017) found that an adequate availability of soil P might
favor the translocation and assimilation of K in tubers.
In Suprema, poor soils were beneficial for tuberization and a higher yield, while soils of
high fertility favored an allocation of N and K to stems and leaves (Figure 2-2, Figure 2-3).
A reduction in translocation of photoassimilates from leaves could be due to the excess of
N, since N inhibits tuberization and increases flow of photoassimilates to new shoots
instead of promoting tuber growth; in our study, this phenomenon might have been due to
the excess of nutrients, mainly N and K. Similar responses to luxury uptake of N in potato
were referenced by Bowen et al. (1999) in cv. Yungay (Group Andigena) in Peru,
Abdallah et al. (2016) in cv. Bintje (Group Chilotanum), and Fontes et al. (2016) in cv.
Atlantic in Brazil. On the other hand, the effects of a luxury uptake of K were discussed by
Kang et al. (2014) and Svenson and Aide (2017) in cv. Atlantic.
A significant interaction crop cycle x location x cultivar (p <0.005) was found for HI, with
no effect of fertilizer dose on HI detected at 150-60 dap, probably, due to a nutrient
balance that favored the better assimilation of N, with high adaptation of cultivars to soil
type. The genotypic differences among the cultivars and fertilizer effect on dry biomass
accumulation in tubers were corroborated by Zebarth et al. (2008) for Group Andigena
and by Giletto and Echeverría (2015) for Group Chilotanum.
The coefficients a for Nuptake in Group Andigena reflect a higher N requirement at initial
stages of growth than those reported for Group Chilotanum, with N uptake equal to 42.24W
0.63 0.63
in cv. Shepody, 37.15W in cv. Russet Burbank (Bélanger et al., 2001), and 36.0W
0.63
in cv. Asterix (Andriolo et al., 2006). The coefficients b for Nuptake obtained in our study
(Figure 2-1A, B) were higher on high fertility soils, but lower on poor soils when compared
to the ones reported for Group Chilotanum (Bélanger et al., 2001). Furthermore, on fertile
soils, a very low efficiency in N use by tubers (NHI of 0.18 to 0.25) was evidenced in
Suprema (Figure 2-4A) as compared to low fertility soils with NHI of 0.65. This finding
proves a better adaptation of Suprema to locations of higher altitudes and lower edaphic
offer of mineral nutrients. This indicates that Suprema possess greater sink strength for N
assimilates (possibly, proteins) that promote tuber growth, similar to found by Snyder et
al. (1977) in cvs. Kennebec, Norchip, and Norland of Group Chilotanum.
A differential response of NHI according to genotype x cycle (p <0.05) (Figure 2-4A) on

low fertility soils, with NHI of 0.57 (Capiro) and 0.65 (Suprema), was similar to that
reported for accessions of Group Andigena (Zebarth et al., 2008) and cv. Bannock Russet
of Group Chilotanum (Giletto and Echeverria, 2015), although, in the latter study, cvs.
Innovator and Gem Russet had higher NHI values of 0.78 and 0.7, respectively, because
these genotypes had earlier senescence.
The PHI for Capiro under non-limiting conditions of fertilization was not affected by the
differences in soil and climate conditions among the locations (Figure 2-4B)
demonstrating once again that this genotype had higher phenotypic plasticity in its
ambient adaptation with comparative advantages over Suprema on fertile soils, where
Suprema had a lower PHI of 0.3 to 0.4 due to the excess of N and K in soil (Table 2-1;
Figure 2-4B). The PHI values of 0.3-0.4 were significantly lower when those reported for
Chocontá soils depleted in P (Figure 2-4B) and favored the higher partitioning of
assimilates into aerial parts of the plants. This was, possibly, due to increased
accumulation of proteins that promoted an excessive vegetative growth inhibiting
Andigena)
tuberization as explained by Abdallah et al. (2016). Jenkins and Mahmood (2003) found
that deficiency of P together with an adequate availability of K and N increased
partitioning of dry biomass into the tubers.
An increased uptake of P by a cultivar could be due to a better adaptation of this cultivar

to a soil type and features of root morphology (higher root density) and longer stolons (the
case of Suprema) together with genetic mechanisms improving P absorption from soil.
The latter mechanism might include expression of high affinity P transporters in roots on
P-deprived soils and low affinity P transporters (for Capiro) on soils with higher P
availability. The cellular mechanisms of P absorption were not yet evaluated in these
cultivars, but this phenomenon related to efficiency of potato nutrition with P on poor soils
was approached by Liu et al. (2017). In potato, evaluation of P uptake and PHI are less
common as compared to Nuptake and NHI; therefore, the current research represents one
of the first reported references on these indexes for Group Andigena.
The coefficients a and b for Kuptake in Capiro and Suprema were highly significant
according to R2 (Figure 2-1E, F) surpassing their counterparts in Group Chilotanum with
0.683
Kuptake = 55.4W (Cogo et al., 2006). An exception was seen for coefficient b in Capiro
on low fertility soils, with values similar to those reported for cv. Asterix (Cogo et al.,
2006); however, Capiro was more efficient in the use of K requiring 500 kg K ha -1 to reach
W of 25 Mg ha-1. A high uptake of K by potato is due to the active involvement of K in
carbon fixation by Rubisco, respiration, synthesis of amino acids (Oosterhuis et al., 2014),
and, additionally, unloading of photoassimilates in short- (AKT2 channels) and long-
distance (H+ transporters/sucrose) transport. Also, K positively influences the mobility of
amino acids in phloem (Lemoine et al., 2013). On the other hand, K favors cell turgor and
water balance of aerial parts in potato plants facilitating transport of carbohydrates and
organic acids (Oosterhuis et al., 2014), besides promoting synthesis of starch in tubers
and amino acids in roots; the latest are employed for growth of leaves, stems, and tubers
(Haeder et al., 1973).
2.7.2 Efficiency of translocation and extraction of N, P, and K by

tubers
The EtN was similar among the two studied cultivars due to characteristics of Group
Andigena having a considerable level of proteins in tubers (Andre et al., 2007; Hardigan
et al., 2017). Allocation of nitrogenous forms into tubers was accentuated towards the
period of tuber maturation, since proteins were needed for rapid tuber growth between
tuberization (75 dap) and tuber filling (125 dap); in this period, tubers accumulated 72-
76% of their total N (Figure 2-5; Table 2-4). At the same time, accumulation of N in tubers
decreased during maximum maturation, probably, due to increasing rates of K
accumulation and, jointly, of starch synthesis. A similar phenomenon was observed by
Snyder et al. (1977) with differences obtained among cvs. Kenebec and Norchip (Group
Chilotanum) and by Chung et al. (2014) with no differences observed among cvs.
Shepody, Russet Burbank, and Innovator.
Under non-limiting conditions of fertilization, in locations of high productive potential, EtP

and EtK in Capiro exceeded the ones in Suprema, with significant differences (P <0.05)
observed in the coefficient b, which once again proved the high sink strength of Capiro.
The EtP and EtK close to 77% and 61%, respectively, at tuber maturity indicated a fast
allocation of P and K to the tubers (Figure 2-5) once the tuberization started 75 dap, as
these were fundamental mineral elements for starch synthesis in tubers. Chung et al.
(2014) showed that 30% of the total P in tubers is stored as starch, with concentrations
around 2.3% P in tubers of cv. Innovator and up to 2.4% P in total biomass according to
the critical dilution curve for P (Zamuner et al., 2016); in this cultivar, partitioning of P
might have been similar to that in Capiro, since it was a medium-size plant with tubers of
high sink strength.
The low efficiency of translocation of N, P, and K (10 to 16%) at the start of tuberization
(55-75 dap) (Figure 2-5) did not differ among the cultivars (tubers accumulated 15-26% N,
P, and K of the total extraction) due to differentiation processes, when small tubers had
more meristematic cells than the storage ones. The tissues with high meristematic activity
contain low to zero amounts of storage starch and proteins and contribute to an increment
in dry biomass, while, during tuber growth, parenchyma had higher growth rates with
respect to other tissues and accumulated starch and proteins; these phenomena were
explained by Snyder et al. (1977) and Chung et al. (2014).
Andigena)
The major allocation of mineral elements to the tubers in Capiro occurred from start of
tuberization and flowering to tuber filling and cessation of flowering in a short period of 50
days with accumulation of 56% N, 60% P, and 63% K of their total extraction by tubers.
Suprema presented a similar accumulation for N, but a lower one for P (56%) and K
(54%) at this same period (Table 2-4) due to a stepped tuberization during crop cycle and
greater proliferation of branches and leaves. Suprema increased allocation of mineral
elements to tubers towards the end of the cycle as promoted by an increased mobility of
K, which favored a later filling of tubers and was directly related to accumulation of dry
biomass in tubers (data not shown).
This phenomenon coincides with the distribution of K in the studied cultivars and is proved
by Santos et al. (2010) who showed the highest sink strength for Capiro between 100-125
dap (80-90 g DW biomass week-1) that gradually decreased by 125-150 dap (30 g DW
biomass week-1). However, for Suprema, three phases in biomass accumulation were
detected: (i) the highest sink strength starting from flowering and tuberization 100-110 dap
(80 g DW week-1); (ii) a decrease towards cessation of flowering 125-130 dap (60 g DW
week-1), and (iii) an increase towards maturation and senescence 130-160 dap (95 g DW
week-1) (Santos et al., 2010). This compartmentalization of photoassimilates at the end of
maturation might vary with extended duration of leaf area, such as favored by irrigation
and K fertilization during tuber filling, similar to that reported by Schilling et al. (2016) for
Group Chilotanum.
2.8 Conclusions
The uptake of N, P, and K by potato plants (Group Andigena) was identified per cultivar
and per phenological stage, with 35% more P extracted by tubers in Capiro than in
Suprema and similar extraction of N and K by tubers in both cultivars. The ratios and
allocation dynamics of N, P, and K in plants could serve to suggest a fertilizer formula
adjusted to phenological stages of Capiro and Suprema and being more precise under
fertigation conditions. Fractionation of fertilizer doses from start of tuberization up to a
maximum flowering should be implemented, since actual field practices in the region
include application of total edaphic fertilizers before start of tuberization, thus, limiting
uptake of P and K by the plants.
Acknowledgments
The authors greatly acknowledge the help of the Faculty of Agricultural Sciences, National
University of Colombia for development of this research. We acknowledge valuable
support of Ingeplant SAS and FEDEPAPA for funding and laboratory analysis. The
authors express their gratitude to research assistants Mrs. Paola Torres, Mrs. Liliana
Arevalo, Mr. Elías Silva, and Mrs. Andrea Barragán for help in the field. We also thank the
growers Mr. Walter Guzmán of Biogenética, Mr. Yovanny Pulido and Mr. Ricardo Rojas
for providing logistic service to this research.
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3. Accumulation of N, P, and K in the tubers of
potato (Solanum tuberosum L. spp.
andigena) under contrasting soils of the
Andean region of Colombia1
Acumulación de N, P y K en tubérculos de papa

(Solanum tuberosum L.) spp. andigena bajo suelos
contrastantes en zona andina de Colombia
Manuel Iván Goméz, Stanislav Magnitskiy, Luis Ernesto Rodríguez,

Aquiles E. Darghan
3.1 Abastract
The relationship between tuber growth and demand for NPK in andigena group of potato
(Solanum tuberosum) is poorly documented under conditions of the Andean region of
Colombia. It is necessary to establish a specific nutrient management for high yields to
changes in edaphic-environmental supply and improve the production of Diacol Capiro
and Pastusa Suprema cultivars. Twelve treatments were evaluated at different stages
(75-100-125-150 days after planting ) of tuber growth using a repeated measures design
with three factors: two cultivars (Diacol Capiro and Pastusa Suprema); three locations
with contrasting soils (Subachoque, Facatativá, and Choconta) and two levels of
fertilization, F0 (unfertilized) and F1 (fertilized) of incomplete nature by differential
fertilization by soil type. A positive correlation between fresh weight, dry weight and
extraction of N, P and K (kg ha -1) by the tubers, beside harvest extraction index was
1
Publicado en Agronomía Colombiana 35(1): 59-67, 2017. Doi:
10.15446/agron.colomb.v35n1.61068
determined. Under optimal conditions of fertilization significant differences (P<0.001)

between factor interactions cultivar x phenology x location for accumulation of N and P
were detected, where ‘Pastusa Suprema’ was less demanding in the HEI of N (1,92 kg
Mg-1 harvest) and HEI of P (0.38 kg Mg-1 harvest) than ‘Diacol Capiro’ and was better
adapted to acid soils of low fertility, in contrast the HEI of K between 5.28 and 5.34 kg Mg -
1
harvest did not show differences between cultivars due to the genotypic characteristics
in the accumulation of dry biomass and starch that make them suitable for industrial use,
in addition it was verified that the nutritional extraction depends on the genetic potential
determined by the interaction with the environmental and edaphic supply.
Key words: nutrient extraction, nutritional requirements, macronutrients, Solanaceae.
3.2 Resumen
La relación entre el crecimiento del tubérculo y la acumulación NPK en cultivares de papa
(Solanum tuberosum) de spp. andigena está poco documentada bajo condiciones de la
zona andina colombiana y es necesaria para establecer un manejo específico de la
fertilización en la obtención de altos rendimientos de los cultivares Diacol Capiro y
Pastusa Suprema ante las variaciones de oferta edáfico-ambiental. Se evaluaron 12
tratamientos en diferentes etapas el crecimiento del tubérculo (75-100-125-150 días
después de siembra) mediante un diseño en medidas repetidas con tres factores: dos
cultivares (Diacol Capiro y Pastusa Suprema); tres localidades con suelos contrastantes
(Subachoque, Facatativá y Chocontá) y dos niveles de fertilización, F 0 (no fertilizado) y F1
(fertilizado) de naturaleza incompleta por la fertilización diferencial realizada por tipo de
suelo. Se presentó correlación positiva entre el peso fresco, peso seco y extracción N, P
y K (kg ha-1) del tubérculo además se determinó el índice de extracción de cosecha (IEC).
Bajo condiciones óptimas de fertilización se presentaron diferencias significativas
(P<0,001) entre las interacciones de los factores cultivar x fenología x localidad para la
acumulación de N y P, donde ’Pastusa Suprema’ fue menos exigente en el IEC de N
(1,92 kg Mg-1 cosecha) y el IEC de P (0,43 kg Mg -1 cosecha) que ‘Diacol Capiro’, además
se adaptó mejor a suelos ácidos de baja fertilidad, en contraste el IEC de K entre 5,28 y
5,34 kg Mg-1 cosecha no presentó diferencias entre cultivares debido a las características
genotípicas en la acumulación de biomasa seca y almidón que los hacen adecuados para
3. Accumulation of N, P, and K in the tubers of potato (Solanum tuberosum L. 67
spp. andigena) under contrasting soils of the Andean region of Colombia
uso industrial, además se comprobó que la extracción nutricional depende del potencial
genético determinado por la interacción con la oferta edáfico-ambiental.
Palabras clave: extracción nutricional, requerimientos nutricionales, macronutrientes,

Solanaceae.
3.3 Introduction
The growth of potato plants depends on the environmental conditions and genotype-
environment interaction (Corchuelo, 2005; Cabezas, 2013), where absorption,
translocation, and accumulation of essential nutrients in the tuber allow starch
accumulation (Corchuelo, 2005; Kumar et al., 2013). Potato stays among the crops of the
highest demand for mineral nutrients per kg of dry matter produced (Sierra et al., 2002;
White et al., 2009; Kumar et al., 2013). The andigena potatoes are well adapted to soils of
low fertility, but respond favorably to NPK fertilization achieving high yields (Ríos et al.,
2010; Hernández et al., 2012). According to Grandett and Lora (1979), Villamil (2005),
and Gómez and Torres (2012), the macronutrients of the highest requirement in andigena
potato plants are K, N, and P followed by Mg, S, and Ca.
In potato, N is the element essential for protein synthesis, respiration, and growth of
tubers (Westermann, 2005; Kavvadias et al., 2012). In soil conditions of mountain tropical
regions, N availability to the plants could be affected by low mineralization rate, low
temperatures, clay soils, and low organic matter content (Castro and Gómez, 2013),
although the excess of N may result in luxury consumption and reduce tuberization (Ruza
et al., 2013). Deficiency of N caused a reduction in dry matter, reduced leaf area and
fewer leaflets that provides less light interception and a lower rate of photosynthesis
(Balemi, 2009; Marouani et al., 2015; Li et al., 2016). Phosphorus promotes root growth
(Aguilar-Acuña et al., 2006), rapid formation of tubers and starch synthesis (Perrenoud,
1993). In strongly acid soils with pH lower when 5.5 of the Andean regions with cultivated
potato, P is fixed and forms precipitates with iron and aluminum, which decreases its
availability (Hernández et al., 2012; Castro and Gómez, 2013). Potassium is essential for
translocation of sugars to the tubers and starch synthesis, a fundamental processes in the
tuber growth and filling (Reis and Monnerat, 2000). Low soil supply of K in sandy soils,
excess of Ca (Kavvadias et al., 2012), salinity and clay minerals of 2:1 type (Castro and
Gómez, 2013) limit availability of K to potato crop.
The relationship between tuber growth and demand for macronutrients NPK in andigena
potatoes is scarcely documented, especially as affected by the changes in edaphic-
environmental conditions of Colombian Andean region. This relationship is important to
establish a specific nutrition management and efficient use of nutrients, and optimize
fertilization moments in order to improve productivity and sustainability of cultivars
Pastusa Suprema and Diacol Capiro as economically important for fresh consumption
and industrial use. The aim of the study was to evaluate the accumulation of N, P and K in
two cultivars spp. andigena, under different stages of tuber growth in contrasting soils of
the Cundiboyacense plateau. The second objective of the research was to analyze the
effect of fertilization on the growth and yield of tuber in order to establish an efficient and
specific management of nutrition as dependent on soil type and plant phenology.

The study was done in 2013 and 2014 in soils contrasting in the level of fertility in the
Andean region of Colombia in representative locations in Cundinamarca province
characterized with high yield potential (> 50 t ha-1) of potato and different environmental
and soil fertility conditions (Table 3-1).
Twelve treatments were evaluated using a repeated measures design with three factors:
two cultivars spp. andigena (Diacol Capiro and Pastusa Suprema), three locations
(Subachoque, Facatativá and Chocontá), and two levels of fertilization, F0 (unfertilized)
and F1 (fertilized). The experimental design was of incomplete nature by differential
fertilization by soil type (Table 3-2), with a within-subjects factor associated with four
phenological critical stages tuber growth adapted from Segura et al. (2006). These stages
were stage II, 70-75 days after planting (dap) (formation of secondary stems and
tuberization initiation); stage III, 90-100 dap (bloom, maximum tuberization, and beginning
of tuber filling); stage IV, 120-125 dap (final flowering, tuber filling); stage V, 150-160 dap
(senescence, maximum filling, and maturation of tubers).
Table 3-1. Edaphoclimatic conditions in the locations of the study.
Subachoque Facatativá Chocontá

Climatic conditions ††
Altitude, msnm 2,680 2,520 2,780
North Latitude 4°57’50.1” 4°49’26.9 ” 5°5’30.37”
West Longitude 74°09’28.1” 74°22’29.7” 73°43’2.04”
Annual precipitation (mm) 870 951 1295
Max. temperature (ºC) 18.7 18.1 16.9
Min. temperature (ºC) 6.3 7.0 4.4
Average temperature (ºC) 12.5 12.6 10.6
Soil characteristics Typic Andic Humic
Hapludand Eutrudept Dystrudept
Soil fertility ††† Low High Low
pH 5.16 6.4 5.5
-1
Al (cmolc kg ) 0.59 0.0 0.1
MO (g kg-1) 171.0 166.7 67.7
CEC (cmolc kg-1) 8.31 31.95 9.52
Texture Loamy Loamy Clay loamy
-1
N (g kg ) 8.5 8.3 3.3
P (mg kg-1) 24.8 39.64 18.18
K (cmolc kg-1) 0.1 3.14 0.68
-1
Ca (cmolc kg ) 6.11 24.26 7.20
-1
Mg (cmolc kg ) 1.29 4.34 1.57
S (mg kg-1) 23.0 29.53 11.58
Saturation K (g 100 g-1) 1.2 9.82 7.14
-1
Saturation Ca (g 100 g ) 75 77 75
-1
Saturation Mg (g 100 g ) 15 13.17 16.4
Ca/K 62.52 7.72 10.51
(Ca+Mg)/K 71.1 9.1 12.8
† Physico-chemical characterization of soils in arable layer (0-30 cm) was done according to
methodology IGAC (2006). The soils were classified according to USDA classification system (Soil
Survey Staff, 2010). †† Environmental data were taken and calculated from IDEAM (2012-2013).
††† Chemical fertility potential was assessed in the topsoil 0-30 cm (Castro and Gómez, 2013).
In each locality, divided plot design with three replicates was used, where the main plot
corresponded to the cultivars and subplots corresponded to the two levels of fertilization.
The contribution of mineral nutrients, fertilizer sources and fractionation of fertilization (F1)
levels in each location were done according to Table 3-1. Planting was done in
experimental units of 50 m2 (135 plants/unit) with a row spacing of 1 m and 0.37 m
between the plants, and useful harvest area of 36 m 2 with a density of 27,000 plants/ha.
Table 3-2. Doses of mineral nutrients applied with fertilization in the studied locations.
Nutrient† Subachoque Facatativa Choconta

-1
(kg ha ) (F1s) (F1f) (F1ch)
N 198 171 192
P2O5 374 261 340
K2O 380 180 348
CaO 40 110 45
Mg 55 70 56
S 37 74 80
B 2,8 3,4 1,2
Zn 5,6 5,6 2,4
Mn 7,0 7,0 3,0
Cu 1,4 1,4 0,6
Fe 2,8 2,8 1,2
† Fertilizers applied according to the method of soil-plant balance (Castro and Gómez, 2013) and
fractioned according to historical management in areas, where the high yields have been obtained
(>50 Mg ha-1): N, 60% sowing, 40% dap 45-50; P, 70% sowing, 30% 45-50 dap; K, 30% sowing
and 70% 45-50 dap. The sources of granular fertilizers were: N-P, DAP; K, KCl (0-0-60) potassium
®
sulfate (0-0-50); Ca, calcium nitrate (25% CaO); Mg, kieserite; Nutricomplet , a complex
micronutrient source B, Zn, Cu, Mn and Fe, based on sulfates.
At each stage of tuber growth four plants were taken from each experimental unit and a
destructive analysis of tubers and aerial part was carried out. At each stage, fresh weight
of tubers (FWT) (kg ha-1) was measured. For dry weight of tubers (DWT), 200 g of fresh
plant material were dried to constant weight in oven at 70°C for 72 h and the
concentrations of N, P, and K was determined in tubers (IGAC, 2006). The methods used
for elemental analysis were Micro-Kjeldahl volumetric for total nitrogen (%), calcination at
475 °C colorimetric with molybdate and ammonium vanadate for total phosphorus (g kg -1),
calcination at 475 °C for K (g kg -1). Using the accumulation of macronutrients per dry
weight values the nutritional extraction (kg ha-1) and nutrient (g kg-1) were quantified per
growth stage of tubers adapted from Cabalceta et al. (2005) and Gómez and Torres
(2012). With the above mentioned variables harvest index (HI) and the rate of nutrient
extraction at harvest (EI) (kg of nutrient extracted per t of harvested tuber) were
evaluated.
Principal component analysis was used to perform an exploratory analysis and reduce the
dimensionality of the set of variables. Two components were analyzed with a cumulative
variability highly significant 87.55% as a result of statistical analysis; the weights of the
components were used as new variables for which the multivariate analysis of variance,
MANOVA (Ravindra and Dayanand, 1999) was performed. The Pearson correlation
matrix of qualitative variables was analyzed and the regression curves were fitted;
additionally, the graphs of the interaction of factors that describe the behavior and
distribution of nutrients were compared. SAS statistical software version 2014 and Stat
graphics version 2010 were used for the graphics.
3.5.1 Accumulation of N, P and K in tuber phenology

At the beginning of tuberization, 75 dap, ‘Pastusa Suprema’ took up 51 kg ha-1 N, and 100
kg ha-1 K, while ‘Diacol Capiro’ took up about 30 kg ha-1 N and 84 kg ha-1 K, with similar
uptakes of P (Figure 3-1) in Humic Dystrudepts (Choconta) with increased extraction and
higher yields than other soil evaluated. These significant differences between the cultivars
for N y K (P<0.05) could be attributed to early and stepped tuberization in ‘Pastusa
Suprema’ and to highly bearing phenotype of this cultivar.
These plant features, apparently, promoted higher demand for assimilate flow into the
tubers at initial growth stages in soils of low fertility, such as of Chocontá. ‘Pastusa
Suprema’ presented better uptake in soils of low fertility Humic Dystrudepts and Typic
Hapludands (Figure 3-1) with reduced availability of N, P, and K (Table 3-1), while
accumulation of ‘Diacol Capiro’ depended directly on increased availability of P in the soil,
where the highest uptake of P was obtained in Andic Eutrudepts slightly acidic soil (Figure
3-1, Table 3-1).
Therefore, it should be performed earlier and gradual fertilization management of

Supreme as related to the fertilization of ‘Diacol Capiro’ that presented accumulation of
NPK 75 dap after the beginning of tuberization (Figure 3-1B, D and F). After this growth
stage, ‘Diacol Capiro’ and ‘Pastusa Suprema’ increased the demand of assimilates that
coincided with increased nutrient transport to tubers and a localized tuberization at this
phenological stage (Figure 3-1). This is consistent with reports Villamil (2005) for ‘Diacol
Capiro’ and comparable with the data found by Sierra et al. (2002) for the cultivar Desiree
in Chile and White et al. (2009) for spp. tuberosum; where the authors reported increase
in NPK accumulation after the beginning of the tuberization. The uptake should be
synchronized with the application of NPK fertilizer to promote initial growth associated
with the growth of stolons, leaves and stems, increased number of tubers and protein
accumulation. In this sense, it is also important to consider factors, such as hormonal
balance, low temperatures and photosynthetically active radiation (Ruza et al., 2013;
Marouani et al., 2015).
In both cultivars, the maximum extraction of N, P, and K by the tubers presented at the
tuber filling stage 150-160 dap, except cv. Diacol Capiro in Subachoque that presented a
maximum extraction at 125 dap (Figure 3-1 B, D and F). Compared with Facatativa and
Choconta, Subachoque likely had more adverse conditions in the availability of N and K
(Table 3-1) and water deficit in this locality, which accelerated ripening and affected
proportionately losses in yield potential because tuberization of ‘Diacol Capiro’ was
grouped into more defined stages 70-100 dap, with greater emphasis on nutrient
availability at flowering stages.
Therefore, it should be avoided stress affecting organs and metabolism sources for filling
of tubers towards the end of the cycle, such as nutritional and hydric deficits, phenomena
that have been reported by Monneveux et al. (2013). Equally, cv. Ica Purace presented
the earliest maximum accumulation of nutrients at 120 dap (Grandett and Lora, 1979), cv.
Pimpernel at 130 dap (Sierra et al., 2002) regarding accumulation at the end of the cycle
for cv. Desiree (Sierra et al., 2002) and cv. Atlantic (Coraspe-León et al., 2009). The
differences in nutrient accumulation were directly related to soil fertility and tuberization
dynamics per cultivar, whether determinate or indeterminate, so it is important to consider

these conditions in fertilizer management.
200 200
SUBACHOQUE (a) (b)
CHOCONTÁ
150 FACATATIVÁ 150
Uptake N (kg ha-1)
100 100
50 50
0 0
0 25 50 75 100 125 150 175 0 25 50 75 100 125 150 175
45 45
(c) (d)
35 35
Ubtake P (kg ha-1)
25 25
15 15
5 5
-5 0 25 50 75 100 125 150 175 -5 0 25 50 75 100 125 150 175
500 500
450 (e) 450 (f)
400 400
Uptake K (kg ha-1)
350 350
300 300
250 250
200 200
150 150
100 100
50 50
0 0
0 25 50 75 100 125 150 175 0 25 50 75 100 125 150 175
dap, Suprema dap, Capiro
Figure 3-1. Accumulation of N P K in tubers during the growing season in andigena

potato, cvs. Suprema (left) and Capiro (right) in Typic Hapludands (Subachoque, ♦);
Humic Dystrudepts (Chocontá, ■) and Andic Eutrudepts (Facatativá, ▲) with fertilization
(fer1) in the altiplano Cundiboyacense. Dap, day after planting. Error bars indicate
standard error.
The P accumulation had significant differences (P≤0.05) between localities and cultivars
mainly for ‘Diacol Capiro’, probably because of its higher nutrient requirement and lower
extraction on soils poor in P (Typic Hapludands and Humic Dystrudepts) (Figure 3-1 D).
Regarding ‘Pastusa Suprema’, it presented a better response in these soils of low fertility
and without differences in extraction in studied soils. Thus, ‘Pastusa Suprema’ showed a
better adaptation of this cultivar to soils low in P and a lower requirement of P applied with
fertilization and probably for its further exploration at the root and mechanisms of
adaptation to acid soils by exudates of root organic acids that improve the availability of P
fixed. These mechanisms of differential absorption have not yet been investigated for
these cultivars.
Phosphorus was the mineral nutrient of less accumulation in tubers as compared to N and
K in evaluated cultivars characterized with 17-40 kg ha-1 uptake P (Figure 3-1 C and D),
which coincided with local study by Villamil (2005) for ‘Diacol Capiro’. The importance of P
should not be underestimated, because its deficiency in potato crops dramatically
reduced the size of tubers, yields (Aguilar-Acuña et al., 2006; Barben et al., 2007;
Hernandez et al., 2012) and tuber quality (Fernandes et al., 2015). Phosphorus deficiency
slowed down the apical growth, resulting in small plants and reduced formation of starch
in tubers, which manifests itself with necrotic spots distributed in the tubers (Hernández et
al., 2012). Several reports indicated that potato plants growing in soils with low P develop
less leaf area, which affected light interception, reduced plant growth and differentiation,
and generated less stolon bearing tubers (Barben et al., 2007). Results obtained by
Balemi (2009) in three genotypes of spp. tuberosum showed that a low supply of P
affected dramatically the plant growth and production of dry matter, also the importance of
P in starch accumulation in potato was comparable with that of K (Perrenoud, 1993).
Accumulation of K in tubers of ‘Diacol Capiro’ was similar in three locations, with a

maximum extraction of 373 kg ha -1 in soils matching high availability of this element in
Choconta and Facatativa (soil saturations with K >7%). ‘Pastusa Suprema’ presented the
maximum extraction of 429 kg ha -1 in Choconta followed by Subachoque (335 kg ha -1)
and Facatativa (232 kg ha-1); this cultivar evidenced a differential extraction (P≤0.05)
curves according to location and that were similar to those of N (Figure 3-1 A and E).
The most uptake rate of K by the tubers of this cultivar was more associated with
adequate environmental supply with lower temperatures <12°C and greater efficiency in
the use of K y N in these soils that allows adequate sink-source management and
potassium fertilization in soils of low fertility balance. In soils of high fertility, such as Andic
Eutrudepts, potato plants could present luxury consumption of N y K, as it was evidenced
in Facatativa, where a similarly extraction was obtained but with greater accumulation of
N and K in the aerial parts and similar yields in Choconta, suggesting lower doses of
these elements in these soil conditions. In addition, the higher average air temperature in
Facatativa over Choconta may cause a higher photosynthetic rate that matches high
growth of the aerial part and could encourage higher activity of enzymes related to
assimilation of K and, therefore, a high demand for this element.
Importantly, ‘Diacol Capiro’ in soils of low fertility of Subachoque with low concentrations
of native soil K (0.1 cmolc kg-1) (Figure 3-1 F) presented an earlier accumulation of K in
the tubers (125 dap) unlike 150 d in other locations (Figure 3-1F). These data contrasted
in this locality with an accumulation of K at the end of the cycle (150-160 dds) for ‘Pastusa
Suprema’, where tuberization and tuber filling was evident by persistent vegetative
development and tuber growth. This implicates a need for more precise handling of
supplementary fertilization in soils of low offer of K. It could be suggested after 100 dap to
keep photosynthetically active and "green" plant until the end of the cycle avoiding early
ripening and to make liquid or foliar fertilization at the late stages of the crop. These
technologies have been evaluated by Horvat et al. (2014) as an alternative in the
supplementary fertilization management of this crop.
The steps of tuber filling and accumulation of K after 100 dap were related directly to the
partition of the dry matter in tubers and were higher than the rest of the plant, which
coincided with high harvest index > 75%, which favors high yield potential. Similar rates of
harvest were found by Sierra et al. (2002) and Sauceda Acosta (2010) for spp. tuberosum
cv. Desireé with yields close to 90 Mg ha-1 and Santos et al. (2010) for cvs. Diacol Capiro
and Pastusa Suprema with yields close to 50 Mg ha -1. In soils Andic Eutrudepts of
Facatativa, the harvest indexes (HI) for Supreme were lower than 55% and lower than
those obtained in low fertility soils with HI >75%, which corroborates a low sink-source
capacity in soils of high fertility associated with high availability of N and K.
3.5.2 Accumulation of N, P, and K among cultivars
The demand for N, P, and K in the cultivars was directly related to the accumulation of
FWT and DWT. A sequence of accumulation of mineral nutrients K> N> P in both
cultivars was observed, similar to that reported by Westermann (2005); Coraspe-León et
al. (2009); Subramanian et al. (2011) and Fallas and Bertsch (2014) in spp. tuberosum,
and by Grandett and Lora (1979), Villamil (2005); Gomez and Torres (2012) and Lefèvre
et al. (2012) in andigena potato.
The significant relationship between nutrient extraction and FWT allowed estimating the
harvest extraction index (HEI) in kg of nutrient extracted per ton of harvestable organ
(Figure 3-1), and is important to determine nutrient requirements in fertilizer
recommendations according to performance targets locals as reported Ciampitti and
García (2008) and Fallas and Bertsch (2014). HEI of N and P showed significant
differences among cultivars with higher yields at 50 Mg ha -1 (P≤0.05) and HEI higher for
‘Pastusa Suprema’ than for ‘Diacol Capiro’ in a positive linear model (Figure 3-1A and B).
For ‘Pastusa Suprema’ HEI was 1.92 kg Mg-1 N, 0.38 kg Mg-1 P; 5.3 kg Mg -1
K, while for
-1 -1 -1
‘Diacol Capiro’ HEI was 2.27 kg Mg N, 0.47 kg Mg P; 5.3 kg Mg K, when compared
with results reported by Sierra et al. (2002) and Ciampitti and Garcia (2008), when in
potato spp. tuberosum HEI was 3.5 kg Mg-1 N, 0.7 kg Mg-1 P; 5.4 kg Mg-1 K with greater
differences for N and P and equal for K.
‘Diacol Capiro’ required 32% more of N and 48% more of P for the yields superior than 50
Mg ha-1 more than ‘Pastusa Suprema’ with significant differences in beta coefficient
according to a positive lineal model in both cultivars (Figure 3-2 A and B). These
variations in N and P uptake in ‘Diacol Capiro’ can be explained by a genotype of low
stature and tuberization more synchronized with better adaptation to fertile soils and
highest genetic potential; in addition this was also associated with accumulation forms
organic an mineral that determine the nutritional characteristics of the tubers. This
confirms the differences in accumulation of NPK between potato genotypes reported by
White et al. (2009), Karam et al. (2009) and Wekesa et al. (2014). It coincides with the
differences in accumulation of these nutrients in cultivars spp. tuberosum made by
Tekalign and Hammes (2005) and White et al. (2009) in identical environments and
andigena cultivars in the Bogota Plateau made by Grandett and Lora (1979).
250 N-Capiro y = 0,0023x - 10,99, (a)

N-Suprema R² = 0,9458**
200
N uptake (kg ha-1)
150
100 y = 0,002x + 0,9887,

R² = 0,80512**
50
0
0 10 20 30 40 50 60 70 80 90
50 y = 0,0005x - 0,6094
P-Capiro R² = 0,84685** (b)
40 P-Suprema
P uptake (kg ha-1)
30
20 y = 0,0004x + 2,7127
R² = 0,85868**
10
0
0 10 20 30 40 50 60 70 80 90
500 y = 0,0053x + 2,9223,

450
K-Capiro R² = 0,95218 ns (c)
400 K-Suprema
K uptake (kg ha-1)
350
300
250
200 y = 0,0052x + 5,6299,
150 R² = 0,9612 ns
100
50
0
0 10 20 30 40 50 60 70 80 90
FWT (Mg ha )
-1
Figure 3-2. Relationship between the uptake in tubers of N (A), P (B) and K (C) (kg ha -1)
yield (FWT), and cvs. Capiro and Suprema in soils of Cundiboyacense plateau. ** The
value of coefficients b were significant at the 0.05 probability level. ns, not significant the
value coefficients b.
Significant differences (P≤0.05) in extracting P in favor of cultivar Diacol Capiro regarding

‘Pastusa Suprema’ (Figure 3-1B) can be explained by the most concentrated tuberization,
the better translocation of assimilates and increased demand of sinks than that in
‘Pastusa Suprema’. This features in ‘Diacol Capiro’ facilitate uniform ripening with a
defined harvest, which requires a greater demand for ATP and may be associated with
organic and inorganic forms of P. These forms of P could serve as a reserve and will
explain responses in the dormancy period of the tubers that lasts up to 4 months in this
cultivar and is longer than that in ‘Pastusa Suprema’ that presented an earlier period of up
to two months.
The differences in P suggest a higher concentration of phytate complex for ‘Diacol Capiro’
than that in ‘Pastusa Suprema’, which should be evaluated in future studies to test this
hypothesis. In relation to P in the tubers Subramanian et al. (2011) mentioned that this
element was metabolized from organic bonds orthophosphate-ester in starch that
accumulate early in tuber formation and from mineral forms of complexes of phytic acid
(myo-inositol hexakisphosphate) towards maturation of tuber and sprouting and is
necessary for proper formation of seeding tubers. The latter complex may reach between
0.1 and 0.27% DWT with differences between cultivars spp. tuberosum according to
Phillippy et al. (2004).
HEI for K showed no significant differences between the cultivars and this was the
element that had the highest correlation (r2 >0.95) with both yield and DWT compared to
N and P (Figure 3-1C). According to the results for K, HEI presented independent effects
between cultivar, locality and fertilization, suggesting that the K extraction in these
cultivars of spp. andigena was mainly depended on the genetic potential of the cultivars.
The later corroborate the direct effect of K in transport, storage, and conversion of
carbohydrates in spp. andigena, therefore, the importance of studied cultivars for the
process of industrial transformation whit high dry matter in tuber that was corroborated in
this study.
The K compared to N and P is the nutrient that had the highest correlation in ‘Diacol
Capiro’ (r2 0.95) and ‘Pastusa Suprema’ (r2 0.99) respect to DWT. The relationship
between K and tuber growth matched the one reported by Coraspe-León et al. (2009),
Karam et al. (2009), Sarkar et al. (2010), Kavvadias et al. (2012) for potato spp.
tuberosum and Villamil (2005) and Pérez (2015) for potato spp. andigena. The
accumulation of K was directly related to accumulation of dry matter (% DWT) and starch
in the tubers (Perrenoud, 1993; Westerman et al., 1994; Reis Jr and Monnerat, 2000) of
19-28% DWT for spp. andigena (Jiménez et al., 2009). DWT % evaluated in this study
(20-27%) was similar to those reported by Jiménez et al. (2009) and Ñústez (2011).
3.6 Conclusions
It is necessary a differential management of mineral nutrition in cultivars according to their
phenology and soil type and considering the differences in nutritional extractions with
higher yields to 40 Mg ha-1 in both cultivar. It is suggested the management ‘Pastusa
Suprema’ fertilization in soils of low fertility applying 30% of total extraction N P K before
stage II, beginning of tuberization (75 dap) in a uptake ratio N:P:K of 10-2-12 N as related
to ‘Capiro Capiro’ requires about 15% of the total extraction NPK in the uptake ratio N: P:
K 10 2-18; which shows a higher uptake ratio of K in early stages, so its best response
was in soils Humic Dystrudepts of Chocontá and Andic Eutrudepts of Facatativá where
saturation K soil was >7%. For tuber filling stages, before stage III (maximum tuberization
and start filling), 70% (‘Pastusa Suprema’) and 85% (‘Diacol Capiro’) of the total NPK
required by the tuber with an 8-1-20 uptake ratio for ‘Pastusa Suprema’ and 12-2-25
uptake ratio for ‘Diacol Capiro’ should be applied. These recommendations can improve
the physiological efficiency of these nutrients by taking into account the timing between
doses, ratio, and age. Actually, it is fractionated in two applications before 50 dap and is
not applied according to the nutrient extraction.
Acknowledgements
The authors express their gratitude to INGEPLANT SAS, FEDEPAPA and UNAL for
funding and technical support of the agronomists of INGEPLANT SAS Paola Torres,
Liliana Arevalo, Elias Silva, and Andrea Barragan. We also thank growers Walter Guzman
- Biogenética, Yovanny Pulido, Ricardo Rojas and Carlos Acero for their support in the
development of research.
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4. Potential yield and efficiency of N and K
uptake in tubers of cvs. Diacol Capiro and
Pastusa Suprema (Solanum tuberosum
subsp. andigena)1
Potencial de rendimiento y eficiencia en la demanda de N

y K en tubérculos de cv. Capiro y Suprema (Solanum
tuberosum subsp. andigena)
Manuel Iván Gómez, Stanislav Magnitskiy, Luis Ernesto Rodríguez
4.1 Abstract
The expression of yield in potato cultivation depends on the genotype-environment
interaction, where edaphic nutrient supply and fertilization play an important role for the
optimum growth and development of the tuber. The total tuber yield (FWt), dry weight of
tubers (DWt), harvest index (HI) and nutrient use efficiency in tubers (NUEt) were
evaluated in Andean region in Colombia under different stages of tuber growth (75-100-
125-150 days after sowing) using two cultivars (Diacol Capiro and Pastusa Suprema),
three locations with contrasting soils (Subachoque, Facatativá and Chocontá) and two
levels of fertilization varied by soil type, F0 (unfertilized) and F1 (fertilized). Humic
Dystrudept soils with fertilization (Chocontá) presented a late tuber filling with increases of
48% and 64% DWt in the cvs. Pastusa Suprema and Diacol Capiro, respectively. In
‘Pastusa Suprema’, the highest production potentials were obtained in fertilized soils of
low fertility with increases of 60.9% DWt and 75% HI, while ‘Diacol Capiro’ is better
adapted to soils of medium to high fertility with increases up to 86.7% FWt with respect to
1
Sometido a la revista Agronomía Colombiana
unfertilized soils and related to higher rates of nutrient recovery efficiency (RFt),
accumulated nutrients per tuber yield (EPt) and better NUEt for N. ‘Pastusa Suprema’
presented EPt and negative RFt with HI<45% and the lower NUE of N and K in high
fertility soils, which represents a null response to fertilization and possible mechanisms of
luxury consumption of the evaluated elements.
Key words: nutrient use efficiency, productive potential, luxury consumption,

macronutrients.
4.2 Resumen
Obtener el máximo rendimiento en el cultivo de papa depende de la interacción genotipo
x ambiente, donde el suministro edáfico de nutrientes y la fertilización desempeñan un
papel importante para el óptimo crecimiento y desarrollo del tubérculo. Se evaluó el
rendimiento total de tubérculos (PFt), el peso seco de tubérculos (PSt), el índice de
cosecha (IC) y la eficiencia de uso de nutrientes en tubérculos (UENt) en la región Andina
de Colombia en diferentes etapas de crecimiento de tubérculos (75-100-125- 150 días
después de la siembra) utilizando dos cultivares (Diacol Capiro y Pastusa Suprema), tres
localidades con suelos contrastantes (Subachoque, Facatativá y Chocontá) y dos niveles
de fertilización variable por tipo de suelo, F0 (no fertilizado) y F1 (fertilizado). Suelos
Humic Dystrudepts con fertilización (Chocontá) presentaron un llenado tardío con un
incremento PSt de 48% y 64% en los cvs. Pastusa Suprema y Diacol Capiro,
respectivamente. En ‘Pastusa Suprema’ se obtuvieron los mayores potenciales de
producción en suelos de baja fertilidad con incrementos en rendimiento del 60,9% y con
IC de 75%, mientras que ‘Diacol Capiro’ se adapta mejor a suelos de media a alta
fertilidad con incrementos hasta 86,7% en rendimiento (PFt) con respecto a suelos no
fertilizados, además se encuentra relacionada con la mayor recuperación de nutrientes
del fertilizante por el tubérculo (RFt) y mayor eficiencia de nutrientes para la producción
de tubérculos (EPt) . ‘Pastusa Suprema’ presentaron EPt y RFt negativo con IC <45% y
la menor UENt de N y de K en suelos de alta fertilidad, lo que representa una respuesta
nula a la fertilización y posibles mecanismos de consumo de lujo de los elementos
evaluados.
4. Potential yield and efficiency of N and K uptake in tubers of cvs. Diacol 87
Capiro and Pastusa Suprema (Solanum tuberosum subsp. andigena)
Palabras clave: uso eficiente de nutrientes, potencial productivo, consumo de lujo,

macronutrientes
4.3 Introduction
For 2016 in Colombia, an estimated production of 2,623,700 t ha -1 was achieved on
126,100 ha, with Cundinamarca and Boyacá provinces generating 76% of potato
production (Riascos, 2016). Diacol Capiro and Pastusa Suprema are considered the
cultivars of high economic importance and represent 80% of the area cultivated in the
country for fresh consumption and industrial processing (Ñústez, 2011).
In the Andean region of Colombia, yields of 50-60 t ha-1 have been reported for ‘Diacol
Capiro’ (Gómez and Torres, 2012) and 40-50 t ha-1 for ‘Pastusa Suprema’ (Pérez, 2015)
but vary considerably by location or soil type, with an increase in the potential yield in the
last decade due to the technological improvements in certified seed management,
irrigation, mechanization, and balanced fertilization.
In these cultivars, high application of nitrogen and potassium fertilizers facilitates

significant yields (Ríos et al., 2010; Saravia et al., 2016), but there is low absorption
efficiency of mineral nutrients by the plants, probably due to the shallow root systems
(Poljak et al., 2011). This implies a low nutrient recovery of up to 50% for N (Vos, 2009)
and 70% for K (Gómez and Torres, 2012), a result of the low influx of potassium used by
the roots (Rengel and Damon, 2008). On the other hand, it should be considered that the
availability of water and nutrients in soil (Saravia et al., 2016) as well as the plant genetics
with respect to source-sink efficiency, which is variable for cultivars (Trehan and Singh,
2013), directly affects the efficiency of nutrient transport to the tubers (Poljak et al., 2011;
Giletto and Echeverria, 2015; Fernandes and Soratto, 2016). Nutrient Use Efficency
(NUE) in potato tuber recently has been investigated mainly for N, by Poljak (2011),
Saravia et al. (2016), and Marouani and Harbeoui (2016) and for K by Trehan and Singh
(2013), Wang and Wu (2015); in subsp. tuberosum, while the studies on the efficient use
of nutrients in subsp. andigena are less frequent (Zebarth et al., 2012).
Potato of the subsp. andigena cultivated in the Andean region respond favorably to
fertilization mainly in soils with low nutrient supply (Ríos et al., 2010) and nutrient uptake
rates of about 0.2-0.3 kg m2 are measured (Ríos et al., 2010; Gómez and Torres, 2012),
which can reduce the efficient use of fertilizers and negatively impact the profitability and
environmental sustainability of potato production in the country. The efficiency of nutrient
use for Diacol Capiro and Pastusa Suprema cultivars has not been quantified. The
present research evaluated the production potential, accumulation of dry matter, harvest
index (HI) and NUEt for N and K in tubers in response to the balanced fertilization under
three soils contrasting in their fertility; the study aimed to establish an optimal
management of these mineral nutrients according to the cultivar and soil type.
4.4.1 Location and soils

The experiments were conducted between 2013 and 2014 in three potato producing
locations under contrasting soils of Cundiboyacense plains in the Andean region of
Colombia as shown in the Table 4-1. In each location, a randomized complete block
design was established using an incomplete factorial arrangement in divided subplots-
mixed model with three replicates, where the main plot corresponded to the cultivars
(Capiro and Suprema) and the subplots corresponded to two levels of fertilization (F0 and
F1). The amount of mineral nutrient applied, the sources of fertilizers, and the dose
fractioning for F1 level are shown in Table 4-2.
Twelve combinations among factors were evaluated through a repeated measures design
with three factors: two cultivars (Diacol Capiro and Pastusa Suprema), three locations
(Subachoque, Facatativá, and Chocontá) and two levels of fertilization, Fer0 (unfertilized
plots, initial soil fertility conditions) and F1 (fertilized plots). The design was of an
incomplete nature by the differential fertilization done by soil type (Table 4-2), with an
intra-subject factor in time associated with four phenological stages of tuber growth
adapted from Valbuena et al. (2010): Stage II, 70-75 days after sowing (das) (start of
tuberization); Stage III, 90-100 das (flowering, maximum tuberization and start of tuber
filling); Stage IV, 120-125 das (end of flowering, filling of tuber); stage V, 150-160 das
(maximum tuber filling and ripening). Sowing was done in experimental units of 50 m 2
(135 plants parcela-1), with the distances of 1 m between rows and 0.37 m between the
plants, a useful area of 36 m2 (three central rows) and a density of 27.000 plants ha -1.
The agronomic practices of irrigation, weed and phytosanitary management were carried
out according to the needs of the locations, in such a way that effects of external factors
were minimized. For Subachoque and Chocontá, dolomite type amendments in pre-
sowing were incorporated in soil at rates of 1.5 and 1.0 t ha -1, respectively.
Table 4-1. Environmental and soil fertility characteristics at the study sites.
Environment conditions†† Subachoque Chocontá Facatativá

Altitude, m a.s.l. 2,680 2,780 2,520
North latitude 4°57’50.1” 5°5’30.37” 4°49’26.9”
West longitude 74°09’28.1” 73°43’2.04” 74° 22’29.7”
Annual precipitation, mm 870 1,295 951
Max air temperature, ºC 18.7 16.9 18.1
Min air temperature, ºC 6.3 4.4 7.0
Average air temperature, ºC 12.5 10.6 12.6
Typic Humic Andic
Soil properties †
Hapludand Dystrudept Eutrudept
Soil fertility ††† Low Medium High
pH 4.95 5.5 6.4
Al, cmolc kg-1 0.59 0.1 0.0
Soil organic matter, g kg-1 171 67.7 166.7
CIC, cmolc kg-1 8.3 9.5 31.9
Texture Loam Clay loam Loam
-1
N, g kg 8.5 3.3 8.3
-1
P, mg kg 24.8 28.2 39.6
K, cmolc kg-1 0.1 0.7 3.1
Saturation K, g 100g-1 1.2 7.1 9.8
†Physical-chemical characterization of soils in arable layer (0-30 cm) according to IGAC (2006).
Soils classified according to the USDA classification system (Soil Staff, 2010). ††Environmental
data obtained from IDEAM (2013-2014). †††Potential chemical fertility evaluated in the arable
layer 0-30 cm (Castro and Gómez, 2013).
Table 4-2. Doses of mineral nutrients applied with fertilizers in the study sites.
Nutrient†, kg ha-1 Subachoque Facatativá Chocontá

(F1s) (F1f) (F1ch)
N 198 171 192
P2O5 374 261 340
K2O 380 180 348
CaO 40 110 45
Mg 55 60 56
S 37 74 120
B 2.8 3.4 1.2
Zn 5.6 5.6 2.4
Mn 7.0 7.0 3.0
Cu 1.4 1.4 0.6
Fe 2.8 2.8 1.2
† Recommended fertilization rates derived from the soil-plant balance method (Castro
and Gómez, 2013) and fractioning of fertilizer dose according to historical references in
the areas, where the high yields have been obtained (> 50 t ha-1): N, 60% at sowing and
40% at 45-50 days after sowing (das); P, 70% at sowing and 30% at 45-50 dds; K, 30% at
sowing and 70% at 45-50 dds. Granulated fertilizer sources were: N-P, DAP; K, KCl (0-0-
60), potassium sulfate (0-0-50); Ca, calcium nitrate (25% CaO); Mg, kieserite;
Nutricomplet, complex source of micronutrients B, Zn, Cu, Mn and Fe, based on sulfates.
4.4.2 Plant sampling and analysis

At the four stages of tuber growth, four plants per experimental unit were evaluated.
Destructive analysis of leaves, stems and tubers was done. All parts of the plants were
rinsed with deionized water. For each sampling, fresh leaves, aerial stems and tubers
were weighed separately, a sample of 200 g (fresh weight) in each organ was placed in
paper bag and dried in the oven at 70 °C for 72 h. The dry matter (DW) of each sample
was then weighed and DW accumulation in each organ was evaluated per plant and per
stage. All organs of the dried plants were ground using a 40 stainless steel mesh for the
subsequent chemical analysis. The harvest index (HI) was evaluated at the phenological
stage V, HI = (DWt / DWs) * 100, according to Giletto and Echeverria (2015), where DWt
is tuber dry weight and DWs – shoot dry weight. In dry samples, the nutrient
concentrations in tubers were determined according to the methodology of IGAC (2006).
The amounts of N and K extracted by the tubers were calculated by multiplying the
concentration of the nutrients by the DW accumulated by the tubers at each stage of

growth.
4.4.3 Efficiency use and recovery of mineral nutrients by tubers
Considering the treatments without fertilizer application (F0) with respect to the balanced
fertilization in each location (F1), the indices of NUE were estimated. The N and K
recovery efficiency by the tuber from fertilization or acquisition efficiency (RFt) was
calculated using equation RFt = (Et 1-Et0 / amount of mineral nutrient supplied in the
fertilizer) * 100 (Table 4-2), where Et1 is the nutrient extraction by tubers in fertilized soils
(kg ha-1) and Et0 is the nutrient extraction by tubers in unfertilized soils (kg ha -1). This
equation has been adapted from Fernandes and Soratto (2016). The efficiency use of N
and K by the tubers (NUEt) was estimated as the accumulated dry matter in the tuber /
nutrient accumulation in the tuber, as reported in potato by Poljak et al. (2011) and
Rengel and Damon (2008) for N and K, respectively. In addition, we evaluated the
efficiency of the production of tuber obtained per unit of nutrient accumulated (EPt),
according to the equation adapted from Prochnow et al. (2009): EPt = (FWT1-FWT0) /
(Et1-Et0), where FWT1 is the tuber yield in the fertilized treatment and FWT 0 is the tuber
yield in the control treatment.
4.4.4 Statistical analysis

Multivariate analysis of variance was performed assessing differences in factor interaction
with a confidence level of P<0.01. The Pearson correlation matrix for the qualitative
variables was analyzed and the efficiency indices were adjusted. The statistical program
SAS version 2014 was used and, for the figures, the program Statgraphics version 2010
was employed.
4.5.1 Yield, harvest index, and dry weight of tubers

The DWt and FWt presented highly significant differences (P≤0.01) in response to the
interaction phenology*fertilization (location*cultivar), with a higher yield obtained for
‘Pastusa Suprema’ in fertilized treatments (F1) in Subachoque (70.5 t ha -1) and Chocontá
(73.7 t ha-1) associated with soils of medium to low fertility (Figure 4-1) and related to a
higher accumulation of DWt between 17.1 and 19.98 t ha -1, respectively (Figure 4-2). For
‘Diacol Capiro’, the highest yield was registered in fertilized soils with the higher
availability of K (Table 4-1), Humic Dystrudepts and Andic Eutrudepts in Chocontá (67.3 t
ha-1) and Facatativá (73.1 t ha -1), respectively (Figure 4-1), and directly related to the
accumulation of DWt.
80000 Suprema Capiro
70000
60000
Tuber Yield, FWt (kg ha-1)
50000
40000
30000
20000
10000
0
F0 F1 F0 F1 F0 F1
SUBACHOQUE FACATATIVÁ CHOCONTÁ
Figure 4-1. Yield (FWt) of cvs. Capiro and Suprema at the phenological stage V
(maximum tuber filling and maturation) in the absence of fertilization F0, with respect to
the balanced fertilization by location: F1s, Typic Hapludands (Subachoque); F1ch, Humic
Dystrudepts (Chocontá), and F1f, Andic Eutrudepts (Facatativá). P<0.001 for fertilization
(location * cultivar)
The yields of 70 t ha-1 obtained in both cultivars proved the high genetic potential of these
Andean potatoes under optimal environmental and fertilization conditions; these yields
were dependent on the soil-plant conditions, where a greater number of tubers and a
better translocation of assimilates were promoted, exceeding the yields of 45 t ha -1
observed by Ríos et al. (2010), Ñústez (2011), and Pérez (2015) in the same cultivars.
For fertilized soils in Humic Dystrupets (Chocontá) (Fer1ch), an increase in yield of 36.4%
and 69.9% was observed for ‘Pastusa Suprema’ and ‘Diacol Capiro’, respectively, with
respect to the initial soil conditions. These values were lower than the ones found in Typic
Hapludands in Subachoque but with higher increases in FWt for 60.9% (‘Pastusa
Suprema’) and 87.1% (‘Diacol Capiro’), which shows the importance of fertilization for
both cultivars in low fertility soils (Table 4-1) and coincides with the report by Ríos et al.
(2010), who found positive responses to fertilization of up to 95% YFt in ‘Diacol Capiro’ in
low fertility andisoles in Antioquia province (Colombia). The lowest response to
fertilization was found in Andic Eutrudepts (Facatativá) with 20.1% for ‘Pastusa Suprema’
and a marginal and non-significant increase with respect to the control treatment (Figure
4-1); in addition, there was a 30.3% increase in yield in ‘Diacol Capiro’, which
corroborates a higher availability of nutrients in this soil with respect to other evaluated
soils.
For two cultivars in the three locations, there were observed positive and significant
responses in yield, having a better response to fertilization in ‘Diacol Capiro’ than in
‘Pastusa Suprema’, with ‘Diacol Capiro’ being better adapted to soils with higher fertility.
‘Diacol Capiro’ performed the best way as a response to a higher edaphic K supply, better
base ratio (Ca/K, Ca+Mg/K) (Table 4-1) and balanced fertilization (Figure 4-1). The
differential response of two cultivars in favor of Humic Dystrudepts (Chocontá) could be
explained by the better balance in the edaphic supply of K (0.68 cmol kg -1) with lower P
fixation, higher average contents of soil organic matter and absence of interchangeable
Al+3 (Table 4-1). This was observed with respect to Typic Hapludands (Subachoque) that
are P fixing soils associated with a low mineralization and presence of Al+3, where lower
levels of available N might be present. Additionally, K deficient levels were observed
below the critical levels (0.1 cmol kg -1) reported by Castro and Gómez (2013) for similar
soils.
The cvs. Pastusa Suprema and Diacol Capiro showed a greater conversion of assimilates
in fertilized Humic Dystrudept soils (Chocontá), with 27.03 and 28.2% of DWt at harvest,
respectively, a value higher than the one reported by Ñústez (2011) in these cultivars
(24% DWt). The greater contribution of % DWt favors their industrial use. In contrast, in
high fertility soils of Andic Eutrudepts (Facatativá), lower yields and a lower accumulation
of assimilates were obtained in tubers with 22.01% DWt and 19.8% DWt in ‘Diacol Capiro’
and ‘Pastusa Suprema’, respectively. The best DWt in ‘Pastusa Suprema’ related to lower
fertility soils fertilized both in Subachoque and Chocontá (Figure 4-2 and Figure 4-3) with
total DWt of 634 to 740 g/plant, respectively. The DWt was higher than those reported by
Ñústez et al. (2009) of 450 g/plant for this cultivar at low fertility soils in the Colombian
potato producing zone of Zipaquirá (2580 m a.s.l.).
100.0
90.0
Harvest index ( g/100 g)
80.0
70.0
60.0
50.0
40.0
30.0
20.0
10.0
0.0
Fer0 Fer1s Fer0 Fer1f Fer0 Fer1ch
Suprema 75.0 75.9 42.5 36.7 83.2 83.1
Capiro 84.0 85.0 75.6 75.1 77.5 79.5
Figure 4-2. Harvest index (HI) in cvs. Diacol Capiro and Pastusa Suprema in the absence
of fertilization Fer0, with respect to balanced fertilization by location: F1s, Typic
Hapludands (Subachoque); F1ch, Humic Dystrudepts (Chocontá), and F1f, Andic
Eutrudepts (Facatativá). P<0.001 for location*cultivar.
‘Diacol Capiro’ in low to high fertility soils presented a HI between 75 and 85% when
compared to ‘Pastusa Suprema’ that presented HI between 75 and 83% only on low
fertility soils in the locations of Subachoque and Chocontá, respectively. ‘Pastusa
Suprema’ limited its assimilate partition at high fertility soils in Facatativá with a lower than
45% HI due to its more indeterminate growth habit and a later growth cycle with less
accumulation of DWt (21.2%); the similar results were obtained by Giletto and Echeverria
(2015) for late-cycle and indeterminate-type cultivars, such as Markies Russet.
The lower HI in ‘Pastusa Suprema’ were related to the lower adaptation of the genotype
because, in Facatativá, it was cultivated at marginal altitudes close 2,500 m a.s.l. and
average air temperatures higher than 13 ºC when compared with environmental
parameters defined by Ñústez (2011). Additionally, there was an availability of soil
nutrients (Table 4-1) which may have generated an excess of N which could inhibit
tuberization and tuber growth, confirming what was reported by Ruza et al. (2013). On the
other hand, ‘Diacol Capiro’ was more adapted to high levels of N and K due to its better
ability to partition assimilate to the tuber characterized by its genotype of a determinate
habit and better relation of aerial shoots/tubers. This makes it a more efficient crop;
coinciding with research conducted by Kleinkopf et al. (1981) and Trehan and Singh
(2013) for subsp. tuberosum.
The HI in ‘Diacol Capiro’ and ‘Pastusa Suprema’ did not show significant differences in
response to fertilization but did present differences in the cultivar x location interaction
(Figure 4-2). The non-significance of HI in response to fertilization is consistent with
results obtained by Zelalem et al. (2009) and Burga et al. (2014), who reported that
increasing doses of N and K, respectively, had no influence on this parameter. The above
can be explained more by adaptation mechanisms and characteristics of the genotype
and also coincides with reports made by Rengel and Damon (2008) and Gileto and
Echevarria (2015).
25000
20000
Tuber dry weight, DWt (kg ha-1)
15000
10000
5000
0
Fer0 Fer 1s Fer0 Fer1f Fer0 Fer1ch
Suprema II Suprema III Suprema IV Suprema V
Capiro II Capiro III Capiro IV Capiro V
Figure 4-3. Tuber dry weight (DWt) in cvs. Diacol Capiro and Pastusa Suprema at four
phenological stages (II, start of tuberization; III, maximum tuberization-start of filling; IV,
filling of tuber; V, maximum filling and maturation) in the absence of fertilization F0, with
respect to balanced fertilization by location for soils Typic Hapludands (Subachoque),
F1s; Humic Dystrudepts (Chocontá), F1ch and Andic Eutrudepts (Facatativá), Fer1f;
P<0.001 for fertilization (location * cultivar).
As for the accumulation of DWt for ‘Pastusa Suprema’ at 125 das (stage IV) and for
Capiro at 100 das (stage III), an earlier filling of tubers was observed with the Typic
hapludands (Subachoque) soil than in the other locations. This is probably because there
were grown in soils of lower fertility and low contribution of K (0.1 cmol c kg-1) compared to
Chocontá and Facatativá soils (Figure 4-3, Table 4-1), where tuber filling in both cultivars
was concentrated towards the end of the cycle, possibly, due to the best availability of
nutrients, mainly K, during all the phenological stages, which presents a gradual and
linear extraction of these elements (Figure 4-3). The greater accumulation of DWt at
stages IV and V in evaluated cultivars could be associated with phenological stages of
higher photosynthetic demands and coincided with stages of high translocation and
assimilation partition verifying that reported by Valbuena et al. (2010) for these cultivars.
4.5.2 Efficiency in the nutrient use by tubers
‘Pastusa Suprema’ in Andic Eutrudepts of high fertility (Facatativá) presented a negative

response in the recovery of N (-10.8 RFNt) and K (-9.6 RFKt) of the balanced fertilizer
applied (Figure 4-4) and a negative physiological response of N (-267.9 EPt N) and K (-
250 EPt K) for tuber production when compared to positive indexes of NUE for ‘Diacol
Capiro’ (Figure 4-5) indicating significant differences between location and cultivar. The
highest RFt of N and K of the applied fertilizer was observed for ‘Diacol Capiro’ in
Facatativá in high fertility soils in basin zone, with RFNt of 70.2% and RFKt of 79.8%;
these indexes were higher than those found in Chocontá with RFNt of 44.8% and RFKt of
42.3% and in Subachoque (38,1% RFNt y 46,9% RFKt) probably because they are soils
that present greater losses of these nutrients by runoff to be located in mountain
landscapes. The location of Chocontá and Subachoque also presented high yield
potential to ‘Diacol Capiro’ and lower than 60% RFNt reported by Vos (2009) for nitrogen
applications between 150-200 kg ha-1 and had similar indices in the acquisition of K as
reported by Gómez and Torres (2012) in cv. Diacol Capiro.
100
90 CAPIRO
80
70 SUPREMA
RFt(kg DWt/ kg Nutrient)
60
50
40
30
20
10
0
-10
-20
-30
-40
-50
RFNt RFKt RFNt RFKt RFNt RFKt
SUBACHOQUE FACATATIVA CHOCONTA
CAPIRO 38.1 46.9 70.2 79.8 44.8 42.3
SUPREMA 45.8 35.0 -10.8 -9.6 34.2 28.1
Figure 4-4. Efficiency of recovery of N and K in tubers (RFt), kg nutrient extracted per 100
kg nutrient applied in balanced fertilization, cvs. Diacol Capiro and Pastusa Suprema on
contrasting soils of the Andean region-Colombia. P<0.001 in N and K for location *
cultivar.
500.0
400.0
300.0
EPt(kg / kg Nutriente)
200.0
100.0
0.0
CAPIRO
-100.0 SUPREMA
-200.0
-300.0
-400.0
EPt N EPt K EPt N EPt K EPt N EPt K
CAPIRO 365.6 174.4 200.2 172.8 347.1 361.2
SUPREMA 291.1 238.3 -267.9 -250.0 307.9 283.9
Figure 4-5. EPt of N and K (kg of tuber harvested per kg of nutrient extracted) in ‘Diacol
Capiro’ and ‘Pastusa Suprema’ in contrasting soils of the Andean region-Colombia.
P<0.001 in N and K for location * cultivar.
‘Pastusa Suprema’ presented low recovery efficiency (<45%) of nitrogen (RFNt) and
potassium (RFKt) with lower rates than ‘Diacol Capiro’ for K in low fertility soils (Figure
4-4). This is a characteristic of high yield and indeterminate cultivars as reported by
Kleinkopf et al. (1981) for cvs. Russet Burbank and Centennial Russet. In addition, the
low K efficiency was, probably, due to a low supply of unexchangeable K to the roots with
low K extraction in these soils associated with a low root/shoot ratio. Similar results were
reported by Trehan and Singh (2013) for the cultivars subsp. tuberosum: Kufri Jyoti and
Kufri Badshah. Therefore, it is necessary for soils with low availability of K to increase the
diffusion and mass flow of this element by means of an adequate contribution and
fractionation of K in the mineral and/or organic fertilizers starting from sowing. This helps
to counteract the effect of antagonistic elements, such as Al +3, maintaining a balance in
the Ca+2 and Mg+2 ratio, thus, improving the rhizosphere environment and root growth. In
Humic Dystrudepts soils of lower fertility in Chocontá, where the best yields were
obtained, a better physiological efficiency in the production of tubers by nutrient extracted
(EPt) was observed, with significant differences in the interaction between the location
and the cultivar (Figure 4-5). Under these conditions, ‘Diacol Capiro’ and ‘Pastusa
Suprema’ produced EPt of N 347 and 308 kg FWt / kg N, respectively, and EPtK of 361
and 283 kg FWt/ kg K. These were higher results for N and similar for K to those reported
by Trehan and Singh (2013), with EPtN between 250 and 318 kg FWt/kg N and EPtK
between 256-360 kg FWt/kg K for efficient subsp. tuberosum cultivars Kufri and a hybrid
JX 576.
The above relates to the higher response in FWt and DWt from ‘Diacol Capiro’ and
‘Pastusa Suprema’ to fertilization in this location and coincides with the greater Use
Efficiency of N and K in the tubers (Figure 4-6), which can be explained by the better
partition and conversion of the N, K and assimilates to the tubers with a better sink
strength of the tubers at the times of filling for both cultivars under the edaphic-
environmental conditions of this location. In addition, the greater efficiency in cv. Diacol
Capiro under contrasting soils coincides with the better adaptation of this cultivar in a
wider range of altitudes (1,800-3,200 m) and soils, similar to optimal environmental
conditions reported by Ñústez (2011).
‘Diacol Capiro’ responded better to fertile soils, which coincides with HI> 75% (Figure 4-2)
and where lower soil loss factors were observed due to the location in lacustrine basins
areas and the high native K and N content (Table 4-1). Again, this confirms that the best
adaptation for ‘Diacol Capiro’ is in flat areas and in the high fertility soild of the Plateau of
Bogota. This provides a better efficiency in the translocation of nutrients which are
assimilated to the tuber and a high removal of N and K from the plant (Figure 4-4), which
needs to be replenished in fertilization plans. The better adaptation of this cultivar can be
explained by possible differential absorption mechanisms with a greater flow of K and N to
the root and into the tuber. In addition, the presence of specialized channels that also
favor the assimilate translocation together with a lower ratio of aerial shoots/tuber are
mechanisms that have been explained for K by Trehan and Singh (2013) and Wang and
Wu (2015) and for N by Vos (2009) that verified the differences between the genotypes.
Under conditions of high availability of K (3.14 cmol c kg-1) in Andic Eutrudepts (Facatativá)
‘Pastusa Suprema’ responded negatively to the fertilization of this nutrient with EPt K of -
250 and presented the lowest K EPt for ‘Diacol Capiro’ of 172.8 (Figure 4-5). Additionally,
the tuber production response for ‘Pastusa Suprema’ was marginal for high potassium
saturation soils (> 9%) in Andic Eutrudepts (Facatativá) as shown in Figure 4-1. The
above data suggest a possible luxury uptake for K for ‘Pastusa Suprema’ and marginal
uptake for ‘Diacol Capiro’, the phenomena that have been explained for subsp. tuberosum
by Kang et al. (2014). On the other hand, Karam et al. (2009) for cvs. Derby and Umatilla
Russet found less efficiency in the use of K with K 2O levels higher than 289 kg ha-1. In
addition, Burga et al. (2014) verified in tretaploid cultivars that high levels of K affected the
development of tubers to the detriment or marginal of yield, where the excess of this
element might limit the transport of other assimilates or hormones.
On the other hand, Facatativá soils had the lowest physiological efficiency of N (NUEt of
18 kg kg-1 for ‘Diacol Capiro’ and 36 kg kg-1 for ‘Pastusa Suprema’) and K (NUEt of 36 kg
kg-1 ‘Pastusa Suprema’ and 30 kg kg-1 ‘Diacol Capiro’) with contributions of 171 kg ha -1 of
N and 180 kg ha-1 of K, respectively, under high availability of N and K (Table 4-2). This
coincides with findings by Zebarth et al. (2012) and Saravia et al. (2016), who found for
potato low values in the NUE between 40 and 10 kg kg -1 respectively increasing the
availability of N in crops with doses between 200 and 300 kg ha -1 of N can help with
positive plant responses.
140.0 CAPIRO SUPREMA

120.0
NUEt(kg DWt/ kg Nutrient)
100.0
80.0
60.0
40.0
20.0
0.0
NUEt N NUEt K NUEt N NUEt K NUEt N NUEt K
CAPIRO 73.2 37.2 36.0 30.5 97.3 106.7
SUPREMA 49.3 40.1 18.2 18.2 26.8 29.8
Figure 4-6. Efficient use of N and K in tubers of balanced fertilization, NUEt (kg of dry
matter of the tuber per kg of nutrient extracted) in cvs. Diacol Capiro and Pastusa
Suprema in Typic Hapludands, Subachoque, Andic Eutrudepts, Facatativá, and Humic
Dystrudepts, Chocontá in Andean region-Colombia.
The highest physiological efficiency in the use of nutrients in tubers was obtained in
‘Diacol Capiro’ with NUEt of 97.3 kg kg -1 for N and 106.7 kg kg-1 for K in fertilized soils of
the lower fertility in Choconta. This corroborates the better efficiency of ‘Diacol Capiro’ as
a characteristic of the genotype already mentioned and verifies that the best NUE is
significantly different to that in ‘Pastusa Suprema’. ‘Diacol Capiro’ appears to be a
genotype of a determinate type that agrees with results by Kleinkopf et al. (1981) for
cultivars of similar type. These indices for soils of lower fertility were superior than those
reported by Poljak et al. (2011) with NUEt of 71 to 76 kg kg-1 for N in subsp. tuberosum
and N fertilizer contribution between 150-200 kg ha-1, probably, due to the positive
interaction of nutrients in the balanced fertilization of macro and micronutrients in
unsaturated soils with an improvement in the efficient use of nutrients in tropical soils
corroborating that discussed by Prochow et al. (2009).
The low physiological efficiency of N and K in the tubers for cv. Pastusa Suprema in soils
with excesses of N might suggest a luxury consumption of these elements that could be
due to a lower growth of tubers by the low transport of assimilates to organs associated
with low HI (Figure 4-2). With a higher average temperature than in the other locations,
this could result in a high ratio of aerial shoots/tubers, thus, limiting the flow of carbon and
nutrients to tubers similar to that reported by Fandika (2012) and Saravia et al. (2016) for
N and Wang and Wu (2015) for K. These authors suggested agronomic and genetic
strategies to decrease the air shoot/tuber ratio and to improve the translocation of
assimilates into the tubers. According to Roumeliotis (2012) excesses of N and high
temperatures limit tuber formation and growth. Conversely, Fandika (2012) also reported
a less efficient use of N by lower partition of assimilates into the tubers, with a low number
of physiological sources at the start of tuberization under high doses of N.
The imbalance due to excess of N available in soil during tuber filling at the start of
tuberization could also cause a reversal of tubers into stolons affecting the productive
potential, a phenomenon that should be evaluated in these cultivars in future research.
Similar results were proposed by Güller (2009) in potato subsp. tuberosum cv. Ana,
where it was found that at doses higher than 200 kg ha-1, a smaller number of tubers were
present at the tuberization stage with higher generation of source structures. In addition,
Zelalem et al. (2009) reported that doses higher than 140 kg N ha -1 increased the number
of stems and delayed flowering that according to Roumeliotis (2012) affected the
synthesis and transport of the FT-like protein tuberigen that favors tuberization in this
species.
The lower physiological indexes found in the use of K obtained in ‘Pastusa Suprema’ in
high fertility soils (Figure 4-4, Figure 4-5 and Figure 4-6) coincided with a rapid vegetative
growth, a later tuberization and a lower growth of the source organ. This coincides with
the lower accumulation of K and dry matter and the presence of smaller tubers despite
the presence of the same number of tubers as in ‘Diacol Capiro’. This can be explained
through an excess of K, which can generate an imbalance by allowing nitrate
accumulation with a lower assimilation in the aerial part and a decrease in the transport of
carbohydrates and proteins towards the tubers. Similar effects by excesses of K were
found by Kang et al. (2014).
4.6 Conclusions
‘Diacol Capiro’ is more efficient in the use of N and K than ‘Pastusa Suprema’
independent of the soil type and location, although ‘Pastusa Suprema’ presented the best
responses and physiological indexes in soils of lower fertility in higher altitude and lower
ambient temperature with response to balanced fertilization. This suggests the use of
integrated management of fertilization aimed to improve the availability of N and K in the
rhizosphere (acidity and nutrient balance) and a specific fertilization in both cultivars,
considering the environmental supply per site and the soil supply given by soil
pedogenesis.
Acknowledgments
The authors express their gratitude to Ingeplant SAS, Fedepapa and Universidad
Nacional de Colombia for funding and technical support of the agronomists of Ingeplant
SAS Paola Torres, Liliana Arévalo, Elías Silva, and Andrea Barragán. We also thank
growers Walter Guzmán - Biogenética, Yovanny Pulido, Ricardo Rojas and Carlos Acero
for their support in the development of research.
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5. Diagnóstico de K+ y NO3ˉ en savia para
determinar el estado nutricional en papa
(Solanum tuberosum L. subsp. andigena)1
Diagnosis of K+ and NO3ˉ in sap to determine nutritional

status in potato (Solanum tuberosum L. subsp.
andigena)
Manuel I. Gómez-S., Stanislav Magnitskiy, Luis Ernesto Rodríguez
5.1 Resumen
El análisis de savia es una herramienta de diagnóstico nutricional para realizar ajustes
oportunos de fertilización en cultivos hortícolas. El objetivo de este estudio fue determinar
los niveles de referencia de NO3ˉ y K+ en savia por etapa fenológica y conocer el uso
adecuado de esta herramienta de diagnóstico nutricional de N y K en cultivares Diacol
Capiro y Pastusa Suprema en la Sabana de Bogotá a los 55, 75, 100, 125 y 150 días
después de siembra (dds) en respuesta a cuatro niveles de fertilizante (0, 1.450, 1.900 y
2.375 kg ha-1) y su efecto sobre rendimiento, materia seca (PS) e índice de cosecha (IC).
La mayor concentración de K + en savia de tallos para los dos cultivares se presentó en
etapa vegetativa con 4.800 a 5.000 mg L-1, disminuyendo hasta tuberización con 2.725
mg L-1 sin diferencias significativas entre cultivares; contrario al comportamiento de N-
NO3ˉ, donde se presentó diferencias entre cultivares con una máxima concentración en
tuberización con 2.466 mg L-1 en ‘Diacol Capiro’ y 2.200 mg L-1 en ‘Pastusa Suprema’
con disminución en madurez fisiológica. Se obtuvieron niveles de referencia en etapa de
1
Publicado en Revista Colombiana de Ciencias Hortícolas 11(1), 133-142, 2017. Doi:
10.17584/rcch.2017v11i1.6132
floración para ‘Diacol Capiro’ mediante ajuste cuadrático de N-NO3ˉ y K+ en savia con
3.280 mg K L-1 y de 1.231 mg, respectivamente y relacionan con la respuesta a la
fertilización, rendimiento, materia seca y área foliar. En contraste para ‘Pastusa Suprema’
el N evaluado en savia supone un consumo de lujo con niveles superiores a 1,250 mg L-1
de N-NO3ˉ. Con esta técnica diagnóstica en campo se puede establecer ajustes
oportunos en el manejo de la nutrición vegetal nitrogenada y potásica de estos cultivares.
Palabras clave: análisis en savia, indicadores nutricionales, uso eficiente de K y N.
5.2 Abstract
Sap analysis is a nutritional diagnostic tool to make timely fertilization adjustments in
crops. The objective of this study was to determine the reference levels of NO3ˉ and K+ in
sap per phenological stage. The sudy evaluated the proper use of this nutritional
diagnostic tool for N and K in Diacol Capiro and Pastusa Suprema on the Bogota Plateau
at 55, 75, 100, 125 and 150 days after sowing (das) in response to fertilizer levels
balanced (0; 1,450; 1,900 and 2,375 kg ha-1) on the yield, dry matter and harvest index to
determine levels for cultivation and phenology. The highest concentration of K + in sap in
stems occurred in vegetative stage with 4,800-5,000 mg L-1, decreasing in tuberización
with 2,725 mg L-1 , without significant differences between cultivars; contrary to the
behavior of N-NO3ˉ in the stems, where maximum concentration in tuberización was 2,466
mg L-1 ‘Diacol Capiro’ and 2,200 mg L-1 ‘Pastusa Supreme’, with decreased physiological
maturity. The reference levels were obtained at flowering stage for ‘Diacol Capiro’ by
quadratic adjustment of N-NO3ˉ and K+ in sap with 3,280 mg L-1 K and 1,231 mg L -1 N -
NO3ˉ, respectively, and related with the response to fertilization, yield, dry matter, and leaf
area. In contrast for ‘Pastusa Suprema’, the evaluated N, with levels higher than 1,231 mg
L -1 N -NO3ˉ in sap, supposed luxury consumption. Using this diagnostic tool in the field,
adjustments can be made to the management of nitrogen and potassium nutrition of these
cultivars.
Key words: sap analysis, nutrient indicators, efficient use of K and N.

5. Diagnóstico de K+ y NO3ˉ en savia para determinar el estado nutricional en 109
papa (Solanum tuberosum L. subsp. andigena)
5.3 Introducción
La papa (Solanum tuberosum L.) es el tercer producto cultivable para alimento más
importante del mundo (De Jong, 2016). En Colombia constituye uno de los productos
agrícolas de mayor demanda de fertilizantes con el 17 al 20% de los costos totales de
producción para los cultivares Diacol Capiro y Pastusa Suprema usados frecuentemente
para consumo fresco e industrial (Gómez y Torres, 2012), además, es una de las
especies de mayor exigencia nutricional por kg de biomasa seca producida (Kumar et al.,
2013)
Los nutrientes minerales que más extrae la papa bajo condiciones de los Andes son
potasio, (K+) y nitrógeno (N) (Gómez y Torres, 2012). Así el diagnóstico vegetal temprano
de estos elementos esenciales permite ajustar planes de fertilización con el fin de
incrementar producción, reducir costos y disminuir el impacto ambiental (Lefevré et al.,
2012; Vijay et al., 2013).
La savia corresponde a un líquido extraído de tejidos conductores tanto del xilema como
del floema de la planta (Cadahía et al., 2008; Gangaiah et al., 2016). Su uso como
herramienta de análisis nutricional es usado para diagnosticar de manera rápida y
económica deficiencias o excesos de nutrientes (Errebhi et al., 1998; Aquilera et al,
2013). Los reportes para N-NO3ˉ en savia de papa son más frecuentes (Badillo-Tovar et
al., 2001; Moulin et al., 2012; Aguilera et al., 2014) que evaluaciones para K+ (Hochmuth,
1994; Kelling et al., 2002) (Tabla 5-1), debido al mayor impacto ambiental y de
sostenibilidad que representa las pérdidas de N en el ecosistema (Goffart et al., 2008;
Ziadi et al., 2012).
Generalmente, el contenido de NO3ˉ y K+ en savia se realiza en peciolos o tallos de papa

en base seca o fresca porque son estructuras más sensibles a cambios en la
disponibilidad de N y K del suelo o sustrato (Vitosh, 1998; Rogozińska et al., 2005; Moulin
et al., 2012). Adicionalmente, es necesario la calibración de índices por fenología desde
etapas tempranas (Brink et al, 2002; Moulin et al., 2012) para relacionarlos con
componentes de rendimiento y establecer niveles de suficiencia (Badillo-Tovar et al.,
2001; Moreira et al., 2011; Ziadi et al., 2012; Mohr y Tomasiewicz, 2012) con el fin de
realizar ajustes tempranos en los programas de fertilización para optimizar la

productividad del cultivo (Cadahía et al., 2008; Moulin et al., 2012).
Tabla 5-1. Niveles de NO3ˉ y K+ en savia de peciolo de papa en diferentes estados

fenológicos evaluados con medidores de ion selectivo.
ˉ +
Etapa Subespecie NO3 -N savia K savia Referencias
-1 -1
(mg L ) (mg L )
(I) Vegetativa Tuberosum 1.200-1.400 4.500-5.000 Homusht (1994)
Tuberosum 1.400-1.450 Errebhi et al. (1998)
Tuberosum 1.250-1.500 Badillo-Tovar (2001)
(II) Tuberización Tuberosum 1.000-1.400 4.000-4.500 Homusht (1994)
Tuberosum 1.400-2.000 Moreiro et al. (2011)
Tuberosum 1.400-2.750 Badillo-Tovar (2001)
Tuberosum 1.500-2.000 3.000-6.000 Cadahia (2008)
(III) Máxima Floración Tuberosum 1.000-1.200 4.000-4.500 Homusht (1994)
Tuberosum 500-1.000 3.000-4.000 Cadahia (2008)
(IV) Llenado de Tuberosum 900-1.200 3.500-4.000 Homusht (1994)

tubérculo
Tuberosum 850-900 Errebhi et al. (1998)
Tuberosum 900-1.550 Badillo-Tovar (2001)

(V) Maduración
Tuberosum 600-900 2.500-3.000 Homusht (1994)
Andigena 800-900 Aguilera et al. (2014)
Tuberosum 500-600 Errebhi et al. (1998)
Tuberosum 600-850 Badillo-Tovar (2001)
Las diferencias en la acumulación de nutrientes en savia entre cultivares y subespecies

de papa han sido reportados por diferentes autores como se muestra en la Tabla 5-1,
para la mayoría de los casos estos resultados han sido obtenidos por etapa fenológica
como respuesta a diferentes niveles de fertilización o variación por tipo de suelo. Otros
factores que influyen sobre la concentración de nitratos en savia son: (i) hora del día
(Vitosh y Silva, 1996; Rogozińska et al., 2005), (ii) partición y formas de nitrógeno en la
planta (Kolbe, 1997; Mäck y Schjoerring, 2002); (iii) época de siembra (Aguilera et al,
2014), (iv) cultivar y material de propagación (Moreira et al., 2011); (v) disponibilidad de
agua en el suelo y (vi) fuente de fertilizante (Cadahía et al., 2008; Ziadi et al., 2012).
El objetivo de este estudio fue determinar los niveles de referencia de NO3ˉ y K+ en savia
por etapa fenológica y conocer el uso adecuado de esta herramienta de diagnóstico
nutricional para N y K en cultivares Diacol Capiro y Pastusa Suprema cultivados en
suelos de alta fertilidad en la Sabana de Bogotá.
5.4 Materiales y métodos

El estudio se realizó en el año 2015 en el Centro de Investigación en Nutrición en Plantas
del Trópico (CENIPLANT), ubicado en Facatativá al occidente de la Sabana de Bogotá-
Colombia (4° 49’ 26,9” N, 74° 22’ 29,7” O, 2520 msnm). Las condiciones ambientales
presentaron clima frío seco con precipitación anual promedio de 850 mm; temperatura
media máxima de 18,5ºC; temperatura media mínima de 7ºC y temperaturas medias de
12,6 º C y humedad relativa de 85% (IDEAM, 2015). El suelo de estudio se clasificó como
Andic Eutrudept de acuerdo a la Soil Survey Staff (2010). Se caracterizó como un suelo
fluvio lacustres de alta fertilidad y de textura franco arcillosa y altos niveles de N y K
disponible (Tabla 5-2).
Se utilizó un diseño estadístico en medidas repetidas (DMR) con dos factores entre
sujetos: dos cultivares (Diacol Capiro y Pastusa Suprema) y cuatro dosis de fertilizantes
(0, 1.450, 1.900 y 2.375 kg ha-1 de fertilizante) mediante un arreglo en parcelas divididas
con tres réplicas. La serie en el tiempo como factor intra sujetos fue adaptada de
Valbuena et al. (2010) y se asoció a cinco etapas fenológicas críticas de crecimiento del
tubérculo: etapa I (50-55 dds), desarrollo de tallos principales e inicio de crecimiento
vegetativo, etapa II (70-75 dds), formación de tallos secundarios-inicio de tuberización;
etapa III (90-100 dds), floración, máxima tuberización e inicio de llenado; etapa IV (120-
125 dds), final de floración-llenado de tubérculo; etapa V (150-160 dds), senescencia,

máximo llenado y maduración del tubérculo.
Tabla 5-2. Propiedades químicas del suelo en el sitio de evaluación.
Propiedades del suelo Valores
pH 5,82
Al, cmolc kg-1, Metodo Yuang, AA* <0,001
MO, g kg-1, Walkey-Black 12,71
CICE, cmolc kg-1 19,14
N total, g kg-1, Walkey-Black 6,4
P, mg kg-1 , Bray II-Colorimetría 70,16
K, cmolc kg-1, acetato de amonio, AA* 0,87
Ca, cmolc kg-1 , acetato de amonio, AA* 15,95
Mg, cmolc kg-1, acetato de amonio, AA* 2,14
Na, cmolc kg-1, acetato de amonio, AA* 0,18
S, mg kg-1, fosfato monobásico-Colorimetría 30,01
Saturación de K (%) 4,53
Saturación de Ca (%) 87,35
Saturación de Mg (%) 11,18
Ca/K 18,4
(Ca+Mg)/K 20,87
* Determinación por absorción atómica.
Se comparó un tratamiento control con la oferta edáfica del suelo sin aplicación de
fertilizante (Nivel 0) y tres niveles de fertilizante (Tabla 5-3): nivel 1, 75% de la dosis de
fertilizante óptima balanceada; nivel 2, 100 % como dosis de fertilizante óptimo
balanceado y el nivel 3 con el 125 % de la dosis de fertilizante óptima. La recomendación
de fertilización propuesta se realizó teniendo en cuenta el balance en la relación suelo-
planta. Se fraccionó el 55% del N en siembra y el 45% a los 45 dds; el 80% del P a la
siembra y el 20% a los 45 dds; el 12% del K a la siembra y el 88% a los 45 dds. Para el
Mg y menores se aplicó el 63% a la siembra y el 37% a los 45 dds; se usaron la
siguientes fuentes granuladas: N-P, DAP (18-46-0), Nitrax-S (28-4-0); K, KCl (0-0-60);
Mg, kieserita (25% MgO), Nutricomplet; B, Zn, Cu, Mn y Fe, fuente compleja de
micronutrientes con base en sulfatos.
Tabla 5-3. Aporte de nutrientes minerales en los tratamientos con fertilización edáfica
(dosis en kg ha-1).
Nutriente Nivel 1 Nivel 2 Nivel 3

1.450 1.900 2.375
N 123 164 205
P2O5 216 288 360
K2O 176 235 294
Mg 60 80 99
S 113 150 188
B 1,7 2,3 2,9
Zn 3,5 4,6 5,8
Mn 4,2 5,6 7,0
La siembra se realizó en parcelas de 50 m2 (135 plantas), con una distancia entre surcos
de 1,00 y 0,37 m entre plantas, un área útil de cosecha de 36 m2 con un área
experimental de 1.200 m2, para una densidad de 27.000 plantas/ha. Las prácticas
culturales de riego, manejo de arvenses y manejo fitosanitario se realizaron de manera
uniforme y comparables con el manejo comercial. Para cada etapa fenológica y unidad
experimental se evaluó cinco plantas mediante análisis destructivo en hojas, tallos y
tubérculos materia fresca (Pf), materia seca (Ps). El Ps se determinó pesando muestras
de 200 g de material vegetal fresco a peso constante en una estufa de secado a 70 °C
durante 72 horas, adaptado de Moreira et al. (2011).
Se extrajo en campo el jugo celular de los tallos y de los tubérculos en las diferentes
etapas fenológicas para cinco plantas por unidad experimental, se tomó una alícuota de
extracto de savia de 0,5 ml en tres tallos principales de la quinta a sexta hoja verdadera
del ápice y se evaluaron cuatro tubérculos por planta de calidad “primera” con diametro
entre 6-9 cm. La medición del N-NO3ˉ y K+ en savia se realizó de forma directa en campo
entre las 8 y 10 am (Vitosh y Silva, 1996), usando el método de electrodo selectivo de
iones-ISE evaluado por Goffart et al. (2008), Carson et al. (2016) mediante equipos
portátiles Horiba LAQUA twin (Horiba Europe, Leichlingen, Alemania) Ión-K+ y Ión N-
NO3ˉ.
Los datos reportados fueron valores promedios de tres réplicas con el error estándar de
la media. Los datos de las variables fueron analizados mediante análisis de varianza
donde se evaluaron las interacciones de los factores fenología x dosis de fertilizante x
cultivares, y para las medias de las variables se aplicó la prueba de comparación múltiple
de Tukey (P<0,005), además, se ajustaron las curvas de regresión utilizando el programa
estadístico INFOSTAT versión 2014.
5.5 Resultados y discusión
5.5.1 K+ y N-NO3ˉ en savia por etapa fenológica y cultivar

Se encontraron diferencias altamente significativas (P<0,0002) en la concentración de K+
en savia de tallos en ‘Diacol Capiro’ con 4.750 mg L-1, mientras para ‘Pastusa Suprema’
fue de 5.037 mg L-1, siendo mayor en etapas iniciales a los 55 dds y para los tratamientos
fertilizados con interacciones entre el cultivar y fenología tanto para K + (P<0,0004) como
para NO3ˉ (P<0,0001). ‘Diacol Capiro’ mostró una mayor sensibilidad a las variaciones de
K+ (Figura 5-1A) en comparación a ‘Pastusa Suprema’ (Figura 5-1B), esta última no
presentó diferencias significativas de K+ en savia al incremento de la fertilización debido
probablemente a la baja traslocación a los órganos vertederos y menor requerimientos de
fertilización potásica en suelos fértiles. Las concentraciones en etapas iniciales fueron
similares a las reportadas por Hochmut et al. (1994) y Rosen et al. (1996) e inferiores a
6.000 mg L-1 en savia fresca de peciolo, si se compara con lo reportado por Cadahia
(2008) para la subsp. tuberosum.
0Kg/ha 1450Kg/ha
5500 1900Kg/ha 2375Kg/ha 5500
K+ savia de tallo (mg L -1) a
5000 5000 a
4500 4500 a
a
b a a
4000 4000
a
3500 a 3500 a
3000 a a
a 3000 a
b a
2500 a 2500 a
b a
2000 2000
50 75 100 125 150 50 75 100 125 150
Días después de la siembra A Días después de la siembra B
Figura 5-1. Variación de K+ en savia fresca de tallo medida en campo en el ciclo del
para cada etapa fenológica presenta diferencias estadísticas significativas (Tukey,
P<0,05).
A partir de los 55 dds para ‘Diacol Capiro’ y ‘Pastusa Suprema’ hasta los 100 dds en
floración se observó una disminución drástica en las concentraciones de K + en savia de
tallo con valores de 2.316 y 2.516 mg L-1, respectivamente. Disminución en las
concentraciones de K se pueden relacionar con procesos de translocación el inicio de la
etapa de llenado, máxima tuberización y floración; periodos importantes para el
diagnóstico nutricional de estos cultivares, coincidiendo con las etapas fenológicas
críticas para la toma de análisis de savia en papa como lo referencia Hochmut et al.
(1994) y Cadahía (2008). Las mayores concentraciones de K + en savia en las primeras
etapas del cultivo se debe probablemente a una acumulación inicial de asimilados en
estructuras vegetativas de hojas y tallos, necesarias para el crecimiento e inicio de
tuberización, donde la concentración de K + fue menor a medida que se desarrolla el
cultivo, posiblemente como consecuencia de un efecto de dilución por crecimiento y
transporte a órganos vertederos confirmando lo planteado por Kelling et al. (2002).
‘Diacol Capiro’ incrementó la concentración de K+ en savia desde los 100 dds hasta los
150 dds alcanzando valores de 3.300 mg L-1 con diferencias significativas por variación
en la dosis del fertilizante (Figura 5-1A), mientras para ‘Pastusa Suprema’ la
concentración de K+ fue similar y tiende a incrementarse a partir de los 125 dds hasta los
150 dds lo cual relaciona con el llenado más tardío dada a su tuberización continua y una
acumulación de K+ más al final del ciclo respecto a ‘Diacol Capiro’ que presenta una
tuberización más temprana debido a su hábito de crecimiento determinado. ‘Pastusa
Suprema’ no evidenció cambios de K + en savia respecto a niveles de fertilización (Figura
5-1B) probablemente por una menor eficiencia y baja traslocación con alta disponibilidad
de N y K (Tabla 5-2) asociado posiblemente a un crecimiento vegetativo indeterminado
limitando la tuberización y el crecimiento del tubérculo.
El incremento en la acumulación de K+ en la etapa de maduración en ambos cultivares

coincide con el máximo crecimiento del tubérculo, mayor acumulación materia seca y
conversión de almidón, los cuales como órganos vertederos demandan mayor
translocación de asimilados de la parte aérea; contrario al comportamiento de algunos
cultivares de subsp. tuberosum que presentan ciclo más corto y menos tubérculos,
favoreciendo un mayor crecimiento de los vertederos y una disminución gradual de K en
savia de peciolos con niveles hasta de 2.500 mg L-1 en maduración como lo reportó
Homusht (1994). La dinámica en la concentración de N- NO3ˉ en savia de tallo por
fenología en ambos cultivares fue contrario al K+, incrementándose desde los 55 dds en
etapa vegetativa con 1.812 mg L-1 en ‘Diacol Capiro’ y con diferencias significativas por
variación de los niveles de fertilización (Figura 5-2A), mientras en ‘Pastusa Suprema’ los
N-NO3ˉ aumentó desde 1.450 mg L- sin cambios por efecto de la fertilización (Figura
5-2B).
3000 0Kg/ha 1450Kg/ha
3000 0Kg/ha 1450Kg/ha
1900Kg/ha 2375Kg/ha 1900Kg/ha 2375Kg/ha
a 2500
N-NO3 savia tallo (mg L-1)
a a
2500 a a a
a ab 2000
2000 a a a
a b a
a
ab 1500 a
1500 a a
c a
b 1000
1000
c
500 500
0 0
50 75 100 125 150 50 75 100 125 150
Días después de la siembra A Días después de la siembra B
Figura 5-2. Variación de NO3ˉ en savia fresca de tallo medida en campo en el ciclo de
para cada época presenta diferencias estadísticas significativas, Tukey, P<0,05.
A los 100 dds se presentaron los máximos niveles de nitratos en savia de 2.467 mg L-1 y
2.200 mg L-1 (Figura 5-3A) y relaciona con los menores valores de K en savia para ‘Diacol
Capiro’ (Figura 5-3B) y ‘Pastusa Suprema’, respectivamente (Figura 5-2B), este máximo
crecimiento vegetativo coincide con el inicio de la etapa de llenado en la etapa fenológica
III de floración, máxima tuberización, y correlaciona con la mayor area foliar alcanzada
(19.690 cm2) para ‘Diacol Capiro’ que presentó una alta significancia en este modelo
(Figura 5-3), similar a lo encontrado por Valbuena et al. (2010) para el cv. Diacol Capiro
donde el área foliar aumentó rápidamente desde la emergencia y alcanzó su punto
máximo a los 100 dds.
Después de los 100 y 125 dds para ‘Diacol Capiro’ y ‘Pastusa Suprema’, respectivamente
se presentó un descenso en los niveles de nitratos hasta 1.185 mg L-1 para ‘Diacol
Capiro’ por efecto de la traslocación y hasta 1.433 mg L-1 para ‘Pastusa Suprema’ (Figura
5-2) que coincide con un mayor crecimiento vegetativo por su hábito indeterminado en
condiciones de alta disponibilidad de NO 3ˉ debido posiblemente a la reducción y
asimilación primaria del N en aminoacidos. Mäck y Schjoerring (2002) demostraron en
subsp. tuberosum que la actividad en la reducción del nitrato vía nitrato reductasa (NR) y
asimilación del NH4+ vía glutamato sintasa (GS) se presenta mayormente en tallos y
favorece un mayor acumulación de proteínas con crecimiento vegetativo en detrimento
de la crecimiento y desarrollo de tubérculos.
Las menores concentraciones de N- NO3ˉ en savia de tallo para los cultivares evaluados
en etapas iniciales (Figura 5-2) pueden sugerir acumulación de formas orgánicas de N
necesarias para el crecimiento vegetativo de hojas y tallos, diferenciación de estolones a
tubérculos o partición de N a sitios de crecimiento como nuevos brotes. La disminución
de nitratos y conversión a formas proteicas a medida que incrementa el crecimiento del
tubérculo en etapas finales ha sido explicado por Kolbe (1997), Goffart et al. (2008) y
Ruza et al. (2013).
‘Diacol Capiro’ presentó en etapa de máximo llenado y maduración diferencias

significativas como respuesta a los niveles de fertilización con disminuciones drásticas en
las concentraciones de N en savia hasta 800 mg L-1 (sin fertilización) y hasta 1.200 mg L-
1
(1900 kg ha-1 de fertilizante), debido al efecto de la fertilización sobre el incremento en
PS y PF de la planta, lo cual puede también generar un fenómeno de dilución o efectos
por la partición de formas de N hacia tubérculos (Figura 5-3). Este comportamiento fue
similar al encontrado en savia de peciolos por Vitosh (1998) y por Badillo y Tovar (2001)
en etapas de llenado para subsp. tuberosum y por Aguilera et al. (2014) en subsp.
andigena con rangos entre 900 y 1.500 mg L-1 y entre 800-900 mg L-1, respectivamente.
3000 A
2500
NO3- en savia (mg L-1)
2000
1500
y = -5E-06x2 + 0,1969x + 505,99
1000 R² = 0,82989**
500
0
0 5000 10000 15000 20000 25000 30000
6000
B
5000
K+ en savia (mg L-1)
4000
3000
2000
y = 1E-05x2 - 0,4587x + 7310,7
R² = 0,61298*
1000
0
0 5000 10000 15000 20000 25000 30000
Área foliar, AF (cm 2)
Figura 5-3. Relación entre área foliar y el contenido de NO3ˉ (A) y K+ (B) en savia fresca
de tallo medida en campo hasta etapa III (90-100 dds), floración, máxima tuberización e
inicio de llenado en papa ‘Diacol Capiro’ en respuesta a la fertilización en suelos de alta
fertilidad de la Sabana de Bogotá. ** Modelo altamente significativo (P<0,01); * Modelo
significativo (P<0,05).
La mejor respuesta de ‘Diacol Capiro’ a la fertilización puede mostrar una adecuada

eficiencia en la conversión y uso del N que en ‘Pastusa Suprema’, porque ‘Diacol Capiro’
a medida que aumentó la acumulación de asimilados disminuyó los niveles de N
inorgánico en savia y fue la que presentó un mayor crecimiento de tubérculo, lo cual
genera el efecto de dilución y mayor traslocación de K. ‘Pastusa Suprema’ no presentó
cambios del N en savia cuando se incrementó el nivel de fertilización con mayor
acumulación de nitrato en la parte aérea, probamente por la menor demanda nutricional
de N y excesos de N disponible en el suelo, lo que puede favorecer un consumo de lujo
inhibiendo la movilidad del K al tubérculo por un excesivo crecimiento vegetativo en
detrimento de un menor formación y llenado de tubérculos.
Diferencias entre cultivares y fenología en la concentración de N en savia fueron

reportados también por Waterer (1997) y Ziadi et al. (2012). Waterer (1997) reportó que
la concentración de N-NO3ˉ en cuatro cultivares de papa fue menor hacia el final de ciclo
en comparación con plantas jóvenes, sin embargo en la etapas fenológicas iníciales los
cultivares se comportaron de manera diferente, siendo los cultivares Nordona Ranger y
Norkotah los que presentaron un incremento a inicio de floración, en cambio el cv. Alpha
tiende a disminuir la concentración de N-NO3ˉ desde etapas vegetativas.
Mantener bajas las concentraciones N-NO3ˉ hacia el final de ciclo es necesario debido a
que un exceso de nitrógeno, provoca una disminución de la materia seca en tubérculos
por la baja potencia fuente vertedero que puede restringir la acumulación de
carbohidratos en tubérculo y disminuir la calidad para uso industrial. Los problemas
fisiológicos por acumulación de N en la planta han sido discutidos ampliamente por Mäck
y Schjoerring (2002), Ziadi et al. (2012) y Ruza et al. (2013).
El efecto del fertilizante desde etapas tempranas y la sensibilidad en el cambio de N o K

en savia mediante el método de “Cardy meter” encontrado en este estudio fue
comparable a lo reportado por Rosen et al. (1996) y por Mohr y Tomasiewicz (2012) para
el cv. Russet Burbank subsp. tuberosum y por Aguilera et al. (2014) en el cv. Waycha en
subsp. andigena, además ha correlacionado el método Cardy Meter con el de análisis de
tejido en otras especies como en Brassica rapa (Gangaiah et al., 2016) y en tomate por
Carson et al. (2016). Lo anterior convierte a este método de campo en una herramienta
importante para conocer la eficiencia de la fertilización en el cultivo y realizar ajustes

tempranos de estos nutrientes en etapa vegetativa.
5.5.2 N-NO3ˉ y K+ savia y su relación con peso seco y rendimiento

Se presentó una correlación positiva entre el peso seco y fresco de los tubérculos
(r2=0,98 para ‘Diacol Capiro’ y r2=0,95 para ‘Pastusa Suprema’) y la concentración de
NO3ˉ y K+ en savia de tallos, por lo cual, para fines de pronóstico se analizó únicamente
el rendimiento. Por otro lado, evaluaciones de K+ y N-NO3ˉ en savia de tubérculos no
presentó una relación significativa con respecto al Pft para ambos cultivares, lo cual
muestra que para este estudio el órgano fuente no fue el mejor indicador de las
condiciones nutricionales para realizar ajustes de la fertilización debido probablemente a
que el tubérculo no es sensible a cambios en fertilización porque la parte aérea presenta
mayor dinámica metabólica de K y N en procesos fotosíntesis, respiración y actividad
enzimática, posiblemente esto explica la relación directa del N y K en savia de tallo con el
rendimiento que presentó ‘Diacol Capiro’ (Figura 5-4A), coincidiendo con reportes hechos
por Vitosh et al. (1998).
En ‘Diacol Capiro’ el K+ en savia de tallo para los diferentes tratamientos presentó un

modelo cuadrático positivo (r2 =0,77) respecto al rendimiento y un nivel óptimo de K+ de
3.280 mg L-1 con un rango de suficiencia de 3.000 a 3.300 mg L-1 en la etapa de llenado,
el cual relaciona con los máximos rendimientos (Figura 5-4A). Lo anterior sugiere que el
K+ en savia de tallo puede ayudar a diagnosticar en campo el estado nutricional de este
elemento y proyectar la producción de manera oportuna mediante ajustes en el manejo
de la fertilización potásica, cuando se encuentren en rangos de 2.600 a 3.000 mg L-1
(Figura 5-4A). En contraste ‘Pastusa Suprema’ no presentó una variación significativa de
K respecto al rendimiento por la alta disponibilidad de K en estos suelos de alta fertilidad
(Figura 5-4B), de tal manera que se requieren otros estudios en ambientes con un mayor
potencial productivo para este cultivar para determinar niveles óptimos de referencia .
y = -0,0034x2 + 22,328x - 33667, R² = y = -0,0014x2 + 3,4468x + 1281, R² =

4000 0,769 Nivel óptimo de K+ Capiro : 3280
4000 0,97041
3500 mg kg-1 Nivél óptimo de N-NO3- Capiro: 1231 mg
3500 kg-1
Rendimiento (g /planta)
3000 Suprema
3000
2500 Capiro
2500
2000
2000
1500
1500
1000 1000
500 y = -0.0001x2 + 1.3042x - 1608.7 y = 1E+07x-1.271
500
R² = 0.7114 R² = 0.8465
0 0
2250 2500 2750 3000 3250 3500 500 1000 1500 2000 2500 3000
K+ en savia de tallo (mg L-1) A N- NO3- en savia de tallo (mg L-1) B
Figura 5-4. Relación entre las concentraciones K+ (A) y N-NO3ˉ (B) en savia de tallo
medida en campo y el rendimiento, Pft medido desde etapa de floración a maduración
para ‘Diacol Capiro’ (cuadro) y ‘Pastusa Suprema’ (rombo) en suelos de alta fertilidad de
la Sabana de Bogotá.
La relación directa entre K+ en savia y rendimiento se explica por ser el elemento de

mayor extracción que funcionalmente favorece de manera directa el transporte y
conversión se asimilados para el crecimiento del tubérculo, similar a lo reportado por
Gómez y Torres (2012) y Lefèvre et al. (2012) para subsp. andigena en Sabana de
Bogotá y Huancayo-Perú, respectivamente. Estos rangos de suficiencia de K+ en savia de
tallo encontrados en ‘Diacol Capiro’ fueron inferiores a los citados por Homush (1994) y
Kelling et al. (2002) con reportes entre 3.500 a 4.500 mg L-1 de K en savia para el final de
floración y llenado para cultivares de la subsp. tuberosum.
Por otro lado, los contenidos de N- NO3ˉ en savia de tallo disminuyeron a medida que se
incrementaron los Pft en ambos cultivares a partir de un modelo cuadrático negativo
(P<0,001). Los mayores rendimientos se relacionan con un rango entre 800 a 1.231 mg
L-1 de N-NO3ˉ en etapa de llenado, valores superiores a este rango, disminuyen de
manera importante el crecimiento del tubérculo sugiriendo una toxicidad por exceso de
nitrato en detrimento del rendimiento (Figura 5-4B), coincidiendo con lo reportado por
Brink et al (2002) para subsp. tuberosum. Por esto, es necesario mejorar la conversión
de formas inorgánicas a orgánicas que promueven la asimilación de N a proteínas en
estos cultivares de tipo andigena mediante el ajuste de la dosis balanceadas de
nutrientes y así evitar consumos de lujo o toxicidades. Los efectos negativos por
consumos de lujo en papa, exceso de nitratos y su manejo son discutidos por Ziadi et al.
(2012) y Ruza et al. (2013).
Existe una respuesta diferencial entre los cultivares a la fertilización, donde el mayor
potencial de rendimiento lo expresó ‘Diacol Capiro’ con dosis óptima de 1.633 kg ha-1 de
fertilizante y un incremento en rendimiento del 20% (Figura 5-5A), posiblemente por su
mejor eficiencia en suelos de alta disponibilidad de N y K que favorece una mejor
potencia fuente -vertedero; además, de posibles mecanismos para mejor asimilación de
N aún no estudiados en estos cultivares.
Rendimiento, PFt K savia N-NO3 savia

y = -0,0002x2 + 0,6532x + 2896,5
4000 4000 4000 4000
R² = 0,9744; dosis óptima: 1633 kg
Rendimiento Suprema (g planta -1)
Rendimiento Capiro (g planta -1)
iónes en savia de tallo (mg L-1)
3500 3500 3500 3500
iónes en savia de tallo (mg L-1)

3000 3000 3000 3000
y = 0.1665x + 2895.8
2500 2500 2500 y = 0.2332x + 1423.2 2500
R² = 0.9817
R² = 0.9976
2000 2000 2000 2000
1500 1500 1500 1500
1000 1000 1000 1000
500 y = 0.2366x + 715.94 500 500 500
R² = 0.6996
0 0 0 0
0 500 1000 1500 2000 2500 0 500 1000 1500 2000 2500
Dosis de fertilizante (kg ha-1) A Dosis de fertilizante (kg ha-1) B
Figura 5-5. Respuesta en rendimiento en ‘Diacol Capiro’ (A) y ‘Pastusa Suprema’ (B) a la
fertilización y su relación con concentración de K + y N-NO3ˉ en savia de tallo en suelos de
alta fertilidad de la Sabana de Bogotá.
Para ‘Diacol Capiro’ la variación en las dosis de fertilización se relacionó directamente

con el cambio en la concentración de N-NO3ˉ (r2=0,69) y K+ (r2=0,98) con niveles de 3.200
mg L-1 para K+ y de 1.188 mg L-1 para N-NO3ˉ (Figura 5-5A), donde el incremento de
fertilización es proporcional al aumento en la concentración de K+ etapa de llenado e
igual manera el incremento de N-NO3ˉ en savia por efecto de fertilización nitrogenada y
potásica. Lo anterior coincide con resultados presentados por Kelling et al. (2002) y
Rogozińska et al. (2005) para subsp. tuberosum.
En contraste ‘Pastusa Suprema’ presenta una respuesta nula en rendimiento a la

fertilización en suelos de alta fertilidad, pero fue sensible al incremento de nitratos de N-
NO3ˉ hasta 1.400 mg L-1 por aumento de las dosis de fertilizante (r2 = 0, 99) (Figura 5-5B)
que evidencia el consumo de lujo y la acumulación afectando el metabolismo traslocación
y almacenamiento de carbohidratos en el tubérculo en detrimento del potencial de
rendimiento. Dosis altas de N a la fertilización aplicada de 123 hasta 205 kg ha -1,
posiblemente favorece la brotación de tallos secundarios, condición que puede inhibir la
tuberización y disminuir el crecimiento y desarrollo de tubérculos, efectos similares
coinciden con los reportados por Pérez (2015).
Se comprobó en este estudio que los contenidos de nitratos y potasio en savia son
afectados por diversos factores como la fertilización y el tipo de cultivar en suelos de alta
fertilidad, estas variaciones también han sido discutidas ampliamente para esta especie
por Cadahía (2008) y Rogozińska et al. (2005).
5.6 Conclusiones
Se determinó que el análisis de savia en tallos en los cultivares evaluados es una
herramienta de diagnóstico temprano del estatus nutricional y puede usarse como
pronóstico para el manejo de la producción con ajustes en la fertilización principalmente
para ‘Diacol Capiro’ en suelos de alta fertilidad con niveles de referencia óptimos entre
4.500 a 4.700 mg kg-1 para K y entre 1.500-1.700 mg kg-1 para N evaluado desde etapas
vegetativas a los 55 dds, donde niveles superiores a 5.000 mg kg-1 de N y mayores a
1.900 mg kg-1 de K pueden representar consumos de lujo en ambos cultivares. Para el
cv. Diacol Capiro, después de máxima floración, los niveles óptimos en savia que
relacionan con los mayores rendimientos fueron 3.280 mg kg -1 para K+ y 1.281 mg kg-1
para NO3ˉ. Esta técnica en campo puede ser utilizada para identificar aplicaciones
excesivas de fertilizantes nitrogenados, por ello de acuerdo a los niveles encontrados, se
debe mantener una relación K+/NO3ˉ de 2:1 en etapas iniciales y de 3:1 en etapas de
producción para favorecer altos rendimientos con una dosis óptima de fertilizante
balanceado de 1.633 kg para ‘Diacol Capiro’, mientras ‘Pastusa Suprema’ presentó
respuesta nula a la fertilidad en suelos con excesos de N, por ello es importante el
diagnóstico y manejo diferencial de la fertilización por cultivar de acuerdo al hábito de

crecimiento.
Agradecimientos
Los autores expresan su agradecimiento a INGEPLANT SAS por la financiación al
soporte técnico de los Ingenieros Agrónomos Elías Silva y Andrea Barragán. Además
agradecemos al agricultor Ricardo Rojas por su apoyo en el desarrollo de la investigación
5.7 Referencias bibliográficas

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nitrogen concentration measured by ion selective electrode, leaf tissue nitrogen
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meter sap test and icp spectrometry of dry tissue for measuring potassium (K+)
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6. Conclusiones
Se determinaron las relaciones alométricas mediante curvas de dilución de Nc, Pc y Kc
para la subsp. andigena en los cultivares Diacol Capiro y Pastusa Suprema, bajo
condiciones contrastantes de suelos en la zona andina de Colombia. Por primera vez se
obtuvo la curva de dilución crítica de Nc, Pc y Kc para papa de subsp. andigena y es una
de las primeras referencias del Kc para papa.
Las Nc, Pc y Kc que mejor se ajustaron a los cultivares evaluados, Capiro y Suprema,
fueron las que se obtuvieron a partir de la biomasa seca total (W) en comparación a las
obtenidas con el índice de área foliar (IAF) que presentó mayor variación debido a la
rápida translocación de asimilados hacia los tubérculos. Los cultivares evaluados
presentaron diferencias en Kc, donde Capiro presentó un mayor requerimiento en las
etapas iniciales con respecto a Suprema, demostrado en el coeficiente a y con un menor
coeficiente b, lo que muestra una mejor eficiencia en el uso de K para la acumulación de
biomasa seca. Las curvas de dilución crítica fueron similares al modelo propuesto para
subsp. tuberosum, pero con menor dilución en los coeficientes a y b para N y P, lo cual
representa un mayor requerimiento de estos elementos pero con menor eficiencia.
El índice de nutrición (INN) se ajustó a las zonas de estudio mostrando que el N es el

elemento más limitante en la obtención de altos rendimientos principalmente en Capiro,
adicionalmente, se comprobó que Suprema fue más propensa a un consumo de lujo y
que las dosis en suelos de baja fertilidad pueden llegar a un 70% de la dosis óptima de
Capiro. La posibilidad de diagnosticar en etapas tempranas el estado nutricional de N, P
y K, a partir de las curvas de dilución, permite realizar ajustes oportunos en el manejo de
estos elementos para etapas como la tuberización y llenado de turbérculo, y así
potencializar los rendimientos.
Los efectos de la interacción genotipo x localidad para las curvas de consumo de

nutrientes se observó en suelos de baja fertilidad con el cultivar Suprema, donde se
expresa el mayor potencial productivo y una correlación en el índice de cosecha de
nutrientes (ICN, ICP, ICK); mientras que Capiro alcanzo el mayor nivel de adaptación
porque los modelos de consumo de Nr, Pr y Kr fueron significativos en los suelos
estudiados.
El modelo logarítmico positivo fue el que mejor se ajusto a la eficiencia de traslocación

siendo mejor en Capiro con mayor partición de asimilados y nutrientes en el uso de N, P
y K. Además, los requerimientos nutricionales por cultivar y etapa fenológica indicaron un
35% de mayor extracción de P en Capiro que en Suprema y similares para N y K, con
diferenciación en el consumo por etapa fenológica lo cual permite realizar ajustes más
precisos en los planes de fertilización.
Como se ha mencionado y en concordancia con otras investigaciones, Capiro es más

eficiente en el uso de N y K que Suprema independientemente del tipo de suelo, aunque
Suprema presenta las mejores respuestas e índices fisiológicos en suelos de baja
fertilidad a mayor altitud y menor temperatura ambiental con una fertilización balanceada.
De ahí la importancia de un adecuado manejo integrado de la fertilización teniendo en
cuenta la disponibilidad de N y K en la rizosfera (acidez y equilibrio de nutrientes) y una
fertilización específica por cultivar, considerando el suministro ambiental y pedogénetico
del suelo.
El análisis de savia en tallos es una herramienta de diagnóstico temprana del estatus

nutricional y su empleo ayuda a pronósticar el manejo de la producción con correcciones
de fertilización. Los niveles óptimos de referencia para Capiro en suelos de alta fertilidad
son 4.500-4.700 mg kg-1 para K y 1.500 a 1.700 mg kg-1 para N medidos desde etapas
vegetativas de los 55 dds, donde niveles superiores a los 5.000 mg kg-1 de N y 1.900 mg
kg-1 de K representan un riesgo en el consumo de lujo en los cultivares Capiro y
Suprema. En el caso de Capiro, desde la máxima floración los niveles óptimos de 3.280
mg kg-1 para K+ y 1.281 mg kg-1 para NO3- se relacionan con mayores rendimientos.
7. Recomendaciones
Se recomienda en futuras investigaciones relacionar la actividad enzimática junto con la
acumulación de sustancias de tipo fitatos, ésteres de fosfato, almidón sintasa y nitrato
reductasa para conocer aun más la adaptación del cultivar y evaluar su dinámica con los
nutrientes y la calidad del tubérculo.
Con las relaciones NPK y el porcentaje de distribución de estos nutrientes, son el insumo
necesario para el establecimiento de las fórmulas de nutrición en la ferilización
convencional por etapa y puede hacerse de forma precisa en condiciones de fertirriego.
Se sugiere realizar mayor fraccionamiento de la fertilización desde el inicio de la
tuberización hasta la máxima floración ya que actualmente se maneja toda la fertilización
edáfica antes del inicio de la tuberización cuando el potencial de consumo de P y K es
más limitado, principalmente.
Se sugiere para el manejo de la fertilización de Suprema bajo suelos de baja fertilidad, un

aporte del 30% del total extraido de NPK antes de la etapa II cuando inicia la tuberización
(75 dds) en una proporción de 10-2-12. En el caso de Capiro se recomienda un 15% de
la extracción total de NPK en una relación de 10-2-18. En las etapas previas al llenado
antes de la etapa III cuando se presenta la máxima tuberización e inicio de llenado, se
deben fraccionar 70% en Suprema y 85% para Capiro del total NPK requerido con una
relación 8-1-20 para Suprema y 12-2-25 para Capiro, estas recomendaciones pueden
mejorar la eficiencia fisiológica de estos nutrientes al tener en cuenta la sincronización
entre dosis, relación NPK y época. Actualmente se fracciona en dos aplicaciones antes
de los 50 dds y con grados de fertilizante que no se ajustan a la extracción y partición
nutricional.
Las herramientas de diagnóstico nutricional evaluados permiten identificar las

aplicaciones excesivas de fertilizanción nitrogenada. Se recomienda mantener una
relación K+/NO3- de 2:1 para etapas iniciales y 3:1 en etapas de producción con la dosis
balanceada óptima de 1633 kg para Capiro para favorecer altos rendimientos, mientras
Suprema presentó respuesta nula a la fertilidad en suelos con excesos de N, por ello es
importante el diagnóstico y manejo diferencial de la fertilización por cultivar de acuerdo al
hábito de crecimiento.
8. Anexo A: Anovas de rendimiento, eficiencia
de traslocación, índices de cosecha y
covariables
Tabla 8-1. Anovas e interacciones de factores para consumo y eficiencia de traslocación
de N, P y K en cuatro etapas del cultivo e índice de cosecha de nutrientes 150 dds.
Nitrógeno
N.I. 75 dds 100 dds 125 dds 150 dds
CICLO 1 <.0001 <.0001 <.0001 0,0638
LOCALIDAD 1 <.0001 <.0001 <.0001 <.0001
CICLO*LOCALIDAD 2 <.0001 <.0001 <.0001 0,1567
Cultivar 1 0,0229 <.0001 <.0001 0,0009
CICLO*Cultivar 2 0,0029 0,2236 <.0001 0,2813
LOCALIDAD*Cultivar 2 <.0001 0,0099 <.0001 <.0001
CICLO*LOCALI*Cultiva 3 <.0001 0,0138 0,1817 0,9369
Ferti 1 <.0001 0,0364 0,0002 0,8232
CICLO*Ferti 2 0,0011 0,5445 0,4253 0,1735
LOCALIDAD*Ferti 2 0,0098 0,6721 0,0057 0,1289
CICLO*LOCALIDA*Ferti 3 0,0006 0,1009 0,006 0,5006
Cultivar*Ferti 2 0,003 0,7896 0,3048 0,5674
CICLO*Cultivar*Ferti 3 0,0074 0,5489 0,0144 0,508
LOCALI*Cultiva*Ferti 3 0,7726 0,1982 0,0005 0,2923
CICL*LOCA*Cult*Ferti 4 0,0025 0,0464 0,0106 0,9511
Rep 1 0,6542 0,3954 0,709 0,025
Potasio
75 100 125 150
CICLO 1 <.0001 <.0001 <.0001 <.0001
LOCALIDAD 1 <.0001 <.0001 <.0001 <.0001
CICLO*LOCALIDAD 2 <.0001 0,1263 <.0001 0,0619
Cultivar 1 0,0448 <.0001 <.0001 <.0001
CICLO*Cultivar 2 <.0001 0,0253 <.0001 0,3055
LOCALIDAD*Cultivar 2 <.0001 0,0005 <.0001 <.0001
Ferti 1 0,0009 <.0001 <.0001 0,1507
134 Acumulación y distribución de macronutrientes minerales en dos cultivares
de Solanum tuberosum L. en diferentes ambientes del altiplano Cundi-
boyacense
CICLO*Ferti 2 0,0416 0,0544 0,3108 0,7634

LOCALIDAD*Ferti 2 0,0085 <.0001 0,0123 0,779
CICLO*LOCALIDA*Ferti 3 0,0005 0,0252 0,0028 0,8763
Cultivar*Ferti 2 0,4753 0,3298 0,8136 0,1745
CICLO*Cultivar*Ferti 3 0,8763 0,4178 0,3089 0,4289
LOCALI*Cultiva*Ferti 3 0,1714 0,4892 0,0002 0,0329
CICL*LOCA*Cult*Ferti 4 0,0771 0,7278 0,9152 0,8408
Rep 1 0,59 0,3988 0,8434 0,025
Fósforo
75 100 125 150
CICLO 1 <.0001 0,0018 <.0001 0,6279
LOCALIDAD 1 <.0001 <.0001 <.0001 <.0001
CICLO*LOCALIDAD 2 <.0001 0,0006 <.0001 0,0213
Cultivar 1 0,024 <.0001 <.0001 <.0001
CICLO*Cultivar 2 0,0132 0,4404 <.0001 0,1061
LOCALIDAD*Cultivar 2 <.0001 <.0001 <.0001 <.0001
Ferti 1 0,002 <.0001 <.0001 0,0946
CICLO*Ferti 2 0,1321 0,6747 0,0558 0,007
LOCALIDAD*Ferti 2 0,0018 0,3976 0,2288 0,0948
CICLO*LOCALIDA*Ferti 3 0,0001 0,7962 <.0001 0,5846
Cultivar*Ferti 2 0,0985 0,047 0,0046 0,1392
CICLO*Cultivar*Ferti 3 0,0553 0,7155 <.0001 0,1256
LOCALI*Cultiva*Ferti 3 0,3555 0,0088 <.0001 0,1096
CICL*LOCA*Cult*Ferti 4 0,0175 0,1783 <.0001 0,3304
Rep 1 0,6804 0,3942 0,9054 0,0353
Anexo A: Anovas de rendimiento, eficiencia de traslocación, índices de 135
cosecha y covariables
Tabla 8-2. Análisis de regresión de covariables de suelo y clima respecto a las variables
evaluadas.
W ~Env+Gen+Rep%in%Env+Gen*Env
AIC general model

141.759570752765
* Sum Sq Df F value Pr(>F)

Env 37,17057 3 0,79151 0,53193
Gen 16,7688 1 1,07123 0,33094
Env:Rep 141,55946 8 1,13039 0,43331
Env:Gen 340,88844 3 7,25888 0,01136
Residuals 125,23086 8 NA NA
SELECTED MODEL
YLD ~Env+Rep%in%Env+Gen +Gen* CaMgK
AIC selected model

142.34977049551
* Sum Sq Df F value Pr(>F) %GxE %AcumGxE

Env 252,96464 3 5,56115 0,01658 * *
Gen 16,7688 1 1,10593 0,31772 * *
Env:Rep 141,55946 8 1,16701 0,40143 * *
Gen:CaMgK 314,49326 1 20,74138 0,00105 92,25694482 *
Residuals 151,62604 10 NA NA * *
STEPS FOR FACTORIAL

REGRESSION
Effect name Sum Sq Df Fvalue AIC Pr>F TorF
Gen* effect
CaMgK 314,4932617 1 20,74137593 142,3497705 0,001051366 entered
boyacense
GENERAL MODEL
rent ~Env+Gen+Rep%in%Env+Gen*Env
AIC general model 206.535315344362
* Sum Sq Df F value Pr(>F)

Env 1016,78573 3 1,45657 0,29744
Gen 4879,27296 1 20,96899 0,0018
Env:Rep 1890,23919 8 1,01543 0,49163
Env:Gen 5320,34046 3 7,6215 0,00989
Residuals 1861,51925 8 NA NA
SELECTED MODEL
YLD ~Env+Rep%in%Env+Gen +Gen* P
AIC selected model 203.604180435597
* Sum Sq Df F value Pr(>F) %GxE %AcumGxE

Env 3224,51293 3 5,52247 0,01693 * *
Gen 4879,27296 1 25,06951 0,00053 * *
Env:Rep 1890,23919 8 1,214 0,37938 * *
Gen:P 5235,56191 1 26,90011 0,00041 98,40652021 *
Residuals 1946,2978 10 NA NA * *
STEPS FOR FACTORIAL REGRESSION

Effect name Sum Sq Df Fvalue AIC Pr>F TorF
effect
Gen* P 5235,561915 1 26,90010706 203,6041804 0,00040919 entered
Anexo A: Anovas de rendimiento, eficiencia de traslocación, índices de 137
cosecha y covariables
Tabla 8-3. Anovas e interacciones de factores en las variables rendimiento.
Fuente DF Tipo III SS Cuadrado de la media F-Valor Pr > F

CICLO 1 76,5913 76,5913 0,36 0,5479
LOCALIDAD 1 66,63806 66,63806 0,32 0,5751
CICLO*LOCALIDAD 1 2711,10677 2711,10677 12,91 0,0006
Cultivar 1 8535,40918 8535,40918 40,64 <.0001
CICLO*Cultivar 1 7964,45766 7964,45766 37,92 <.0001
LOCALIDAD*Cultivar 1 12019,66028 12019,66028 57,22 <.0001
CICLO*LOCALI*Cultiva 1 25,19876 25,19876 0,12 0,7301
fert 3 5360,78163 1786,92721 8,51 <.0001
CICLO*fert 3 424,14651 141,38217 0,67 0,5714
LOCALIDAD*fert 3 191,96241 63,98747 0,3 0,8219
CICLO*LOCALIDAD*fert 3 91,04808 30,34936 0,14 0,9329
Cultivar*fert 3 1327,75133 442,58378 2,11 0,1072
CICLO*Cultivar*fert 3 405,04707 135,01569 0,64 0,5901
LOCALID*Cultiva*fert 3 1542,52623 514,17541 2,45 0,071
CICL*LOCA*Culti*fert 3 436,32866 145,44289 0,69 0,5598
Rep 3 636,30811 212,1027 1,01 0,3938
Error 69 14493,19906
Tabla 8-4. Anovas e interacciones de factores en las variables rendimiento por cultivar.
Capiro Suprema
Fuente DF F- F-
Tipo I SS CM Valor Pr > F Tipo I SS CM Valor Pr > F
Ciclo 1 2391,11568 2391 28,0 <.0001 4392,9 4393 15,9 0,0003
Localidad 1 7330,94649 7331 85,9 <.0001 4934,4 4934 17,86 0,0002
Ciclo* Localidad 1 1664,69091 1665 19,5 0,0001 1135,8 1136 4,11 0,0508
Fertilización 3 5979,89218 1993 23,3 <.0001 994,7 332 1,2 0,3251
Ciclo*Fertilización 3 434,792268 145 1,7 0,1861 470,7 157 0,57 0,6402
Localidad*Ferti 3 964,308289 321 3,77 0,0197 725,9 242 0,88 0,4636
Ciclo*Loc*Ferti 3 433,263256 144 1,69 0,1874 94,1 31 0,11 0,9516
Bloque 3 883,87331 295 3,46 0,0274 2312,8 771 2,79 0,0558
Error 33 2813,87114 9119,00167
boyacense
Tabla 8-5. Anovas e interacciones de factores para índice de cosecha por cultivar.
Cuadrado
Fuente DF Tipo I SS F-Valor Pr > F
de la media
Ferti 1 0.03073787 0,031 25.93 <.0001
Cultivar 1 0.35995915 0,360 303.64 <.0001
Cultivar*Ferti 1 0.00963250 0,010 8.13 0.0077
LOCALIDAD 1 0.45611488 0,456 384.75 <.0001
Ferti*LOCALIDAD 1 0.00017119 0,000 0.14 0.7065
Cultivar*LOCALIDAD 1 0.30063509 0,301 253.60 <.0001
Cultiv*Ferti*LOCALIDAD 1 0.00007801 0,000 0.07 0.7992
CICLO 1 0.00754653 0,008 6.37 0.0170
Ferti*CICLO 1 0.01833896 0,018 15.47 0.0004
Cultivar*CICLO 1 0.00162872 0,002 1.37 0.2501
Cultivar*Ferti*CICLO 1 0.00125584 0,001 1.06 0.3113
CICLO*LOCALIDAD 1 0.01612147 0,016 13.60 0.0009
Ferti*CICLO*LOCALIDA 1 0.00306552 0,003 2.59 0.1180
Cultiv*CICLO*LOCALID 1 0.02510554 0,025 21.18 <.0001
Cult*Fert*CICL*LOCAL 1 0.00167723 0,002 1.41 0.2433
Rep 3 0.00499937 0,002 1.41 0.2597
9. Anexo B: Coeficientes de consumo,
coeficientes de traslocación e índice de
cosecha de nutriente
Tabla 9-1. Coeficientes de consumo crítico de N, P y K a partir de la biomasa total para
Capiro y Suprema.
Cultivar Suelos Rango W a b IC a 95% IC b 95% SEa SEb R2

Chotá 1-25,6 56,38 0,584 52,52-97,46 0,358-0,585 11,0469 0,0549 0,9033**
Capiro
Faca 1-31,0 73,59 0,658 43,65-128,6 0,457-0,794 20,6712 0,082 0,8246**
N
Chotá 1-28,9 68,13 0,504 62,81-113,8 0,2829-0,497 12,4318 0,0524 0,8682**
Suprema
Faca 1-35,1 78,02 0,692 8,687-89,49 0,6526-1,236 19,4834 8,6872 0,6778*
Chotá 1-25,6 4,267 0,786 4,260-7,810 0,547-0,744 0,8233 0,0484 0,9690**
Capiro
Faca 1-31,0 6,504 0,753 6,970-11,999 0,5259-0,706 1,2276 0,0442 0,9181**
P
Chotá 1-28,9 4,765 0,662 4,8216-11,67 0,2924-0,666 1,678 0,0754 0,8775**
Suprema
Faca 1-35,1 6,724 0,779 4,2468-11,76 0,5665-0,884 1,834 0,0776 0,8942**
Chotá 1-25,6 77,64 0,671 33,505-157,9 0,3877-0,858 30,568 0,1157 0,7595*
Capiro
Faca 1-31,0 79,52 0,854 73,216-186,7 0,4715-0,774 27,659 0,0739 0,8535*
K
Chotá 1-28,9 63,93 0,776 23,211-93,19 0,6252-1,035 17,15 0,1005 0,9103**
Suprema
Faca 1-35,1 42,77 0,772 216.7-40,02 0,1076-1,098 62,67 0,2418 0,3932 ns
IC, intervalo de confianza de los coeficientes al 95%. EE, error estándar de los coeficientes.**
Diferencias significativas p<0,05 de los coeficientes a y b entre los cultivares de estudio; ns, no
existen diferencias significativas. Chtá, suelo en Chocontá de baja fertilidad; Faca, suelo en
Facatativá de alta fertilidad.
boyacense
Tabla 9-2. Coeficientes a y b de eficiencia de traslocación de N, P y K para Capiro y

Suprema sin fertilización (0) y bajo condiciones optimas de fertilización (1).
Intervalo de confianza Intervalo de confianza

Cultivar SEa Seb
a 95% b 95%
Capiro-0 -197.5--22.2311 0.5024-4.2063 20,4 0,430

Suprema-0 -66.4785-54.2487 1.1262-1.3846 1,4 0,030
P
Capiro-1 -121.6--12.7855 0.1762-2.4749 12,6 0,267
Suprema-1 -70.4182-34.8561 0.7124-1.4638 4,1 0,087
Capiro-0 -34.7418-0.0823 -0.0929-0.6429 4,0 0,086
Suprema-0 -74.5129-17.4003 0.3066-1.5134 6,6 0,140
N
Capiro-1 -74.7187-25.1067 -0.6864-1.4229 11,6 0,245
Suprema-1 -35.6707-21.2737 -0.5646-0.6386 6,6 0,140
Capiro-0 -173.8-1.473 -0.0165-3.6859 20,4 0,430
K Suprema-0 -77,025 0.0191-1.6467 9,0 0,189
Capiro-1 -121.8-13.8001 -0.3839-2.4815 15,8 0,333
Suprema-1 -39.6282-18.9836 0.348-0.7842 2,4 0,051
10. Anexo C: Componentes principales y
análisis de diseño en medidas repetidas de
acumulación de biomasa, nutrientes y uso
eficiente de nutrientes enComponente
Gráfica de Pesos del el tubérculo
Fe
0,8
0,6
Componente 3
0,4 Mn N
Ca P
Cu Zn
0,2 B ren
Mg K
0 ps
S Na
-0,2
0,4
-0,4 ps% 0,2
0
-0,2
0 0,05 -0,4 Componente 2
0,1 0,15 0,2 -0,6
0,25 0,3 -0,8
Componente 1
Figura 10-1. Componentes principales de variable y acumulación de nutrientes en

tubérculos.
boyacense
Tabla 10-1. Algoritmo de componentes principales de variables fisiológicas de

crecimiento, consumo y concentración de nutrientes para análisis exploratorio y reducir la
dimensionalidad del conjunto de variables.
OPTIONS LINESIZE=72 NODATE PAGENO=1;
DATA ACP;
INPUT ps psKha rend N P K Mg Ca Na AZ Fe B Cu Mn Zn ;
DATALINES;
15.1 1660.8 10974.0 24.1 3.5 46.5 1.661 0.830 0.415 1.993 0.158 0.022 0.020 0.017 0.031
21.7 5782.2 26600.0 45.7 6.9 137.0 5.204 1.156 1.440 5.782 0.293 0.082
0.023 0.023 0.114
25.8 12174.0 47250.0 108.3 15.8 264.2 12.174 2.435 3.031 15.826 0.722 0.110
0.049 0.090 0.238
25.2 13120.3 52080.0 122.0 17.1 282.1 13.120 2.624 3.267 15.744 0.718 0.102
0.052 0.093 0.220
15.9 839.7 5272.5 7.3 1.4 20.2 0.672 0.420 0.210 1.008 0.034 0.013
0.004 0.007 0.011
23.4 2996.6 12780.0 19.5 3.6 62.0 2.397 0.599 0.746 3.596 0.131 0.012
0.012 0.012 0.030
25.9 5029.5 19440.0 29.7 7.0 102.6 4.024 1.006 1.252 7.041 0.229 0.050
0.020 0.020 0.093
24.7 6268.7 25400.0 41.4 7.5 126.6 5.642 1.254 1.561 7.522 0.316 0.133
0.025 0.025 0.081
18.8 2190.5 11650.0 24.1 5.5 65.1 2.410 1.095 0.548 2.410 0.164 0.033
0.015 0.012 0.043
20.9 6866.2 32900.0 55.6 13.0 155.9 4.806 1.373 1.710 6.180 0.398 0.056
0.027 0.027 0.115
20.6 8100.0 33185.0 65.6 15.4 183.9 5.670 1.620 2.017 7.290 0.469 0.066
0.032 0.032 0.135
24.6 16357.3 66500.0 135.8 27.8 320.6 11.450 4.907 4.073 13.086 1.493 0.274
0.065 0.092 0.272
21.0 1515.6 7230.0 10.5 3.5 39.0 1.364 0.606 0.379 1.516 0.092 0.014
0.008 0.008 0.018
24.3 4952.1 20400.0 19.8 9.4 109.9 3.466 0.990 1.233 4.952 0.208 0.055
0.025 0.020 0.048
26.7 7830.4 29300.0 31.3 14.1 165.2 5.481 1.566 1.950 8.613 0.438 0.059
0.031 0.031 0.108
28.2 11509.2 40860.0 46.0 17.3 203.7 8.056 2.302 2.866 11.509 0.536 0.200
0.046 0.046 0.117
14.6 1661.3 11400.0 29.1 7.1 63.8 2.326 1.495 0.414 2.492 0.301 0.042
0.010 0.025 0.067
15.8 5762.2 36360.0 83.0 19.6 189.0 6.338 1.729 1.435 8.643 0.538 0.118
0.033 0.064 0.192
18.1 8964.2 49560.0 93.2 30.5 278.8 6.275 2.600 2.232 8.964 0.482 0.165
0.047 0.044 0.157
20.0 12765.9 63750.0 159.6 34.5 321.7 12.766 3.702 3.179 14.043 0.772 0.063
0.063 0.083 0.304
13.6 1547.6 11400.0 25.1 6.2 56.6 1.702 0.619 0.385 2.321 0.141 0.020
0.012 0.014 0.047
16.3 3608.9 22100.0 42.6 12.3 118.7 3.970 0.722 0.899 4.692 0.350 0.061
0.027 0.034 0.097
18.1 6820.8 37710.0 70.9 23.2 212.1 6.821 1.978 1.698 8.185 0.720 0.211
0.060 0.062 0.207
20.9 11387.4 54407.1 100.2 30.7 267.6 11.387 3.302 2.835 14.804 0.752 0.056
0.061 0.093 0.219
15.9 668.6 4200.0 10.2 2.9 25.7 0.936 0.669 0.166 1.070 0.083 0.022
0.005 0.010 0.031
18.2 2893.8 15900.0 27.8 6.9 69.5 2.026 1.158 0.721 2.026 0.212 0.051
0.014 0.014 0.049
20.6 6047.6 29400.0 55.6 16.3 148.8 5.443 2.419 1.506 6.048 0.448 0.154
0.040 0.048 0.160
22.9 10889.4 47500.0 102.4 25.0 239.6 8.712 2.178 2.711 9.800 0.551 0.053
0.053 0.053 0.197
16.0 464.8 2902.7 8.4 2.1 16.2 0.604 0.279 0.116 0.837 0.058 0.006
0.005 0.004 0.018
Anexo C: Componentes principales y análisis de diseño en medidas repetidas 143
de acumulación de biomasa, nutrientes y uso eficiente de nutrientes en el
tubérculo
20.3 2496.6 12300.0 24.5 7.5 62.7 2.497 0.999 0.622 2.996 0.212 0.048
0.016 0.018 0.062
23.3 5039.8 21630.0 59.0 16.1 149.2 5.544 1.512 1.255 6.048 0.644 0.137
0.037 0.075 0.157
29.5 11444.7 38838.5 133.9 21.7 240.3 8.011 3.433 2.850 10.300 0.449 0.056
0.056 0.058 0.145
15.8 1892.2 12002.0 27.4 4.0 53.0 1.892 0.946 0.473 2.271 0.180 0.025
0.022 0.020 0.035
20.4 5055.6 24740.0 39.9 6.1 119.8 4.550 1.011 1.259 5.056 0.256 0.072
0.020 0.020 0.099
26.5 12831.9 48500.0 114.2 16.7 278.5 12.832 2.566 3.195 16.681 0.761 0.115
0.051 0.095 0.250
22.0 13101.0 59550.0 121.8 17.0 281.7 13.101 2.620 3.262 15.721 0.717 0.102
0.052 0.093 0.220
15.4 576.6 3735.0 5.0 1.0 13.9 0.461 0.288 0.144 0.692 0.023 0.009
0.003 0.005 0.007
24.4 3523.6 14435.0 22.9 4.2 72.9 2.819 0.705 0.877 4.228 0.154 0.014
0.014 0.014 0.036
25.4 5138.9 20200.0 30.3 7.2 104.8 4.111 1.028 1.280 7.194 0.234 0.051
0.021 0.021 0.095
26.4 7686.8 29100.0 50.7 9.2 155.3 6.918 1.537 1.914 9.224 0.387 0.163
0.031 0.031 0.099
17.5 1979.4 11335.0 21.8 4.9 58.8 2.177 0.990 0.495 2.177 0.148 0.030
0.013 0.011 0.039
20.3 6612.1 32500.0 53.6 12.6 150.1 4.628 1.322 1.646 5.951 0.383 0.054
0.026 0.026 0.110
21.2 8250.0 44600.0 66.8 15.7 187.3 5.775 1.650 2.054 7.425 0.478 0.067
0.033 0.033 0.138
25.4 17730.9 69800.0 147.2 30.1 347.5 12.412 5.319 4.415 14.185 1.618 0.297
0.071 0.100 0.295
20.5 1468.1 7170.0 10.1 3.4 37.7 1.321 0.587 0.367 1.468 0.090 0.013
0.007 0.007 0.017
22.4 5110.6 22800.0 20.4 9.7 113.5 3.577 1.022 1.273 5.111 0.215 0.057
0.026 0.020 0.050
25.2 7246.8 28800.0 29.0 13.0 152.9 5.073 1.449 1.804 7.971 0.405 0.054
0.029 0.029 0.100
28.4 13090.0 46120.0 52.4 19.6 231.7 9.163 2.618 3.259 13.090 0.610 0.228
0.052 0.052 0.133
14.6 1503.9 10320.0 26.3 6.5 57.7 2.105 1.353 0.374 2.256 0.272 0.038
0.009 0.022 0.061
13.7 4664.8 34012.5 67.2 15.9 153.0 5.131 1.399 1.162 6.997 0.435 0.095
0.027 0.052 0.155
17.6 9002.4 51150.0 93.6 30.6 280.0 6.302 2.611 2.242 9.002 0.484 0.166
0.047 0.044 0.158
20.2 16067.3 79462.5 200.8 43.4 404.9 16.067 4.660 4.001 17.674 0.971 0.079
0.079 0.105 0.383
12.1 1164.5 9600.0 18.9 4.7 42.6 1.281 0.466 0.290 1.747 0.106 0.015
0.009 0.010 0.036
15.0 3610.4 24033.3 42.6 12.3 118.8 3.971 0.722 0.899 4.694 0.350 0.061
0.027 0.034 0.097
17.6 6020.3 39150.0 62.6 20.5 187.2 6.020 1.746 1.499 7.224 0.635 0.186
0.053 0.055 0.183
20.3 11873.8 58607.1 104.5 32.1 279.0 11.874 3.443 2.957 15.436 0.784 0.058
0.064 0.097 0.228
15.9 402.4 2535.2 6.1 1.8 15.5 0.563 0.402 0.100 0.644 0.050 0.013
0.003 0.006 0.019
17.7 2862.9 16200.0 27.5 6.9 68.7 2.004 1.145 0.713 2.004 0.209 0.050
0.014 0.014 0.049
20.7 6147.9 29700.0 56.6 16.6 151.2 5.533 2.459 1.531 6.148 0.455 0.156
0.041 0.049 0.162
21.1 9257.8 43975.0 87.0 21.3 203.7 7.406 1.852 2.305 8.332 0.469 0.045
0.045 0.045 0.167
15.9 457.4 2873.3 8.2 2.1 16.0 0.595 0.274 0.114 0.823 0.057 0.006
0.005 0.004 0.018
boyacense
19.4 2420.6 12450.0 23.7 7.3 60.8 2.421 0.968 0.603 2.905 0.205 0.047
0.016 0.018 0.060
23.5 5146.5 21900.0 60.2 16.5 152.3 5.661 1.544 1.281 6.176 0.657 0.140
0.038 0.077 0.160
26.5 9771.8 36840.0 114.3 18.6 205.2 6.840 2.932 2.433 8.795 0.383 0.048
0.048 0.050 0.124
15.8 2054.3 13030.0 29.8 4.3 57.5 2.054 1.027 0.514 2.465 0.195 0.027
0.024 0.021 0.038
22.5 7015.2 31210.0 55.4 8.4 166.3 6.314 1.403 1.747 7.015 0.356 0.100
0.028 0.028 0.138
26.9 13644.9 50800.0 113.0 16.0 262.0 12.200 2.400 3.000 14.400 0.670 0.100
0.055 0.090 0.210
23.0 12217.0 53100.0 121.0 17.7 296.0 13.600 2.700 3.400 17.700 0.810 0.120
0.049 0.100 0.270
15.7 824.3 5250.0 7.2 1.4 19.9 0.659 0.412 0.206 0.989 0.033 0.013
0.004 0.007 0.011
23.5 3190.9 13600.0 20.7 3.8 66.1 2.553 0.638 0.795 3.829 0.140 0.013
0.013 0.013 0.032
26.8 5300.0 19800.0 31.3 7.4 108.1 4.240 1.060 1.320 7.420 0.241 0.053
0.021 0.021 0.098
24.7 8355.4 33800.0 55.1 10.0 168.8 7.520 1.671 2.080 10.026 0.421 0.178
0.033 0.033 0.108
17.5 2142.8 12220.0 23.6 5.4 63.6 2.357 1.071 0.536 2.357 0.161 0.032
0.015 0.012 0.042
22.8 7351.5 32300.0 59.5 14.0 166.9 5.146 1.470 1.831 6.616 0.426 0.060
0.029 0.029 0.123
22.8 8383.2 36700.0 67.9 15.9 190.3 5.868 1.677 2.087 7.545 0.486 0.068
0.034 0.034 0.140
23.0 17334.4 75220.0 143.9 29.5 339.8 12.134 5.200 4.316 13.868 1.582 0.290
0.069 0.098 0.288
20.0 1720.9 8610.0 11.9 4.0 44.2 1.549 0.688 0.430 1.721 0.105 0.015
0.009 0.009 0.020
23.6 4851.4 20520.0 19.4 9.2 107.7 3.396 0.970 1.208 4.851 0.204 0.054
0.025 0.019 0.047
26.8 6809.7 25440.0 27.2 12.3 143.7 4.767 1.362 1.696 7.491 0.381 0.051
0.027 0.027 0.094
29.7 13240.2 44520.0 53.0 19.9 234.4 9.268 2.648 3.297 13.240 0.617 0.230
0.053 0.053 0.135
11.9 1287.2 10860.0 22.5 5.5 49.4 1.802 1.158 0.321 1.931 0.233 0.033
0.008 0.019 0.052
14.4 5020.8 34800.0 72.3 17.1 164.7 5.523 1.506 1.250 7.531 0.469 0.103
0.029 0.056 0.167
16.1 7722.0 48000.0 80.3 26.3 240.2 5.405 2.239 1.923 7.722 0.415 0.142
0.041 0.038 0.136
20.4 15553.7 76225.0 194.4 42.0 392.0 15.554 4.511 3.873 17.109 0.940 0.076
0.076 0.102 0.370
13.6 1425.4 10500.0 23.1 5.7 52.2 1.568 0.570 0.355 2.138 0.130 0.018
0.011 0.013 0.044
15.4 3990.5 25950.0 47.1 13.6 131.3 4.390 0.798 0.994 5.188 0.387 0.067
0.030 0.038 0.107
18.1 6565.8 36300.0 68.3 22.3 204.2 6.566 1.904 1.635 7.879 0.693 0.203
0.058 0.060 0.200
21.2 11841.8 55778.6 104.2 32.0 278.3 11.842 3.434 2.949 15.394 0.782 0.058
0.063 0.097 0.227
15.9 467.2 2943.3 7.1 2.1 18.0 0.654 0.467 0.116 0.748 0.058 0.015
0.003 0.007 0.022
16.9 2863.3 16950.0 27.5 6.9 68.7 2.004 1.145 0.713 2.004 0.210 0.050
0.014 0.014 0.049
20.4 5967.0 29250.0 54.9 16.1 146.8 5.370 2.387 1.486 5.967 0.442 0.152
0.040 0.047 0.157
22.9 11549.8 50425.0 108.6 26.6 254.1 9.240 2.310 2.876 10.395 0.585 0.057
0.057 0.057 0.208
15.4 444.9 2888.0 8.0 2.0 15.5 0.578 0.267 0.111 0.801 0.056 0.006
0.005 0.004 0.017
19.7 2398.4 12150.0 23.5 7.2 60.2 2.398 0.959 0.597 2.878 0.203 0.046
0.016 0.018 0.059
tubérculo
23.4 4998.2 21360.0 58.5 16.0 147.9 5.498 1.499 1.245 5.998 0.638 0.136
0.037 0.075 0.155
24.6 9324.5 37839.2 109.1 17.7 195.8 6.527 2.797 2.322 8.392 0.366 0.046
0.046 0.047 0.119
22.6 2294.1 10168.8 25.2 5.7 68.1 2.523 1.147 0.574 2.523 0.172 0.034
0.016 0.013 0.045
24.6 7843.0 31896.4 74.9 17.3 205.5 7.059 2.745 1.957 7.843 0.521 0.091
0.042 0.037 0.142
25.7 12520.9 48795.6 101.4 23.8 284.2 8.765 2.504 3.118 11.269 0.725 0.102
0.050 0.050 0.209
27.4 21099.2 77100.0 175.1 35.9 413.5 14.769 6.330 5.254 16.879 1.504 0.353
0.084 0.119 0.350
22.4 4574.8 20455.8 31.6 10.5 117.6 4.117 1.830 1.144 4.575 0.279 0.041
0.023 0.023 0.053
25.2 6495.6 25735.4 42.5 14.5 163.2 5.684 2.436 1.623 6.577 0.392 0.057
0.032 0.032 0.077
28.5 8514.9 29841.7 52.6 18.5 209.0 7.238 2.980 2.127 8.728 0.509 0.073
0.040 0.040 0.104
28.8 12337.9 42840.0 80.7 27.6 310.0 10.796 4.627 3.083 12.492 0.745 0.109
0.060 0.060 0.120
23.4 3528.3 15065.2 51.2 7.4 98.8 3.528 1.764 0.882 4.234 1.408 0.046
0.042 0.037 0.065
24.2 7859.8 32438.1 92.0 13.4 195.3 7.860 2.751 1.961 9.825 1.801 0.086
0.062 0.070 0.149
25.6 13245.8 51741.4 117.9 17.2 287.4 13.246 2.649 3.298 17.220 0.785 0.119
0.053 0.098 0.258
28.8 21682.2 75390.0 201.6 23.9 466.2 21.682 4.336 5.399 26.019 1.187 0.169
0.087 0.153 0.364
22.0 4092.0 18597.8 35.6 7.0 98.6 3.274 2.046 1.023 4.910 1.649 0.065
0.020 0.037 0.052
24.4 7697.2 31558.9 56.2 11.9 171.3 6.158 2.694 1.920 10.006 1.726 0.100
0.035 0.050 0.120
26.8 12304.6 45960.2 72.6 17.2 251.0 9.844 2.461 3.064 17.226 0.560 0.122
0.049 0.049 0.226
30.2 19587.6 64965.0 129.3 23.5 395.7 17.629 3.918 4.877 23.505 0.986 0.416
0.078 0.078 0.252
19.9 3872.4 19471.1 42.6 9.7 115.0 4.260 1.936 0.968 4.260 0.290 0.058
0.026 0.021 0.075
22.3 7163.1 32164.7 62.3 10.7 158.3 6.089 1.433 1.784 7.521 0.404 0.057
0.029 0.040 0.120
24.6 10328.4 42037.6 83.7 19.6 234.5 7.230 2.066 2.572 9.296 0.598 0.084
0.041 0.041 0.173
30.0 18095.2 60337.5 150.2 30.8 354.7 12.667 5.429 4.506 14.476 1.290 0.303
0.072 0.102 0.301
22.4 4574.8 20455.8 31.6 10.5 117.6 4.117 1.830 1.144 4.575 0.279 0.041
0.023 0.023 0.053
25.2 6495.6 25735.4 42.5 14.5 163.2 5.684 2.436 1.623 6.577 0.392 0.057
0.032 0.032 0.077
28.5 8514.9 29841.7 52.6 18.5 209.0 7.238 2.980 2.127 8.728 0.509 0.073
0.040 0.040 0.104
28.8 12337.9 42840.0 80.7 27.6 310.0 10.796 4.627 3.083 12.492 0.745 0.109
0.060 0.060 0.120
23.5 4068.2 17341.0 35.4 6.9 98.0 3.255 2.034 1.017 4.882 1.639 0.065
0.020 0.036 0.052
24.3 7443.0 30591.9 54.3 11.5 165.6 5.954 2.605 1.857 9.676 1.669 0.096
0.033 0.048 0.116
26.6 12035.7 45315.0 71.0 16.8 245.5 9.629 2.407 2.997 16.850 0.548 0.120
0.048 0.048 0.221
25.3 24677.8 97695.0 162.9 29.6 498.5 22.210 4.936 6.145 29.613 1.243 0.525
0.099 0.099 0.318
23.4 3528.3 15065.2 51.2 7.4 98.8 3.528 1.764 0.882 4.234 1.408 0.046
0.042 0.037 0.065
24.2 7859.8 32438.1 92.0 13.4 195.3 7.860 2.751 1.961 9.825 1.801 0.086
0.062 0.070 0.149
boyacense
25.6 13245.8 51741.4 117.9 17.2 287.4 13.246 2.649 3.298 17.220 0.785 0.119
0.053 0.098 0.258
28.8 21682.2 75390.0 201.6 23.9 466.2 21.682 4.336 5.399 26.019 1.187 0.169
0.087 0.153 0.364
18.5 2331.9 12617.4 25.7 5.8 69.3 2.565 1.166 0.583 2.565 0.175 0.035
0.016 0.013 0.045
22.1 6562.7 29668.7 48.2 10.2 140.8 5.250 1.313 1.634 6.891 0.355 0.096
0.026 0.026 0.097
27.5 11820.1 42930.8 95.7 22.5 268.3 8.274 2.364 2.943 10.638 0.685 0.096
0.047 0.047 0.198
27.7 17868.8 64500.0 148.3 30.4 350.2 12.508 5.361 4.449 14.295 1.274 0.299
0.071 0.101 0.297
21.1 3816.2 18103.4 26.3 8.8 98.1 3.435 1.526 0.954 3.816 0.233 0.034
0.019 0.019 0.044
23.6 5261.8 22267.5 34.4 11.8 132.2 4.604 1.973 1.315 5.328 0.318 0.046
0.026 0.026 0.063
27.3 6970.9 25506.3 43.0 15.2 171.1 5.925 2.440 1.741 7.145 0.416 0.060
0.033 0.033 0.085
24.9 8295.4 33315.0 54.2 18.6 208.4 7.259 3.111 2.073 8.399 0.501 0.073
0.040 0.040 0.100
26.1 3130.5 11979.8 45.4 6.6 87.7 3.130 1.565 0.783 3.757 1.249 0.041
0.037 0.032 0.058
28.7 8646.2 30178.8 101.2 14.7 214.9 8.646 3.026 2.157 10.808 1.981 0.095
0.069 0.077 0.164
27.5 13883.4 50400.0 123.6 18.0 301.3 13.883 2.777 3.457 18.048 0.823 0.125
0.056 0.103 0.271
25.4 18279.8 72000.0 170.0 20.1 393.0 18.280 3.656 4.552 21.936 1.001 0.142
0.073 0.129 0.307
25.6 2974.4 11599.9 25.9 5.1 71.7 2.379 1.487 0.744 3.569 1.199 0.048
0.015 0.027 0.038
25.7 6957.9 27072.1 50.8 10.8 154.8 5.566 2.435 1.736 9.045 1.560 0.090
0.031 0.045 0.109
27.9 12363.6 44263.4 72.9 17.3 252.2 9.891 2.473 3.079 17.309 0.563 0.123
0.049 0.049 0.227
25.6 16827.2 65775.0 111.1 20.2 339.9 15.145 3.365 4.190 20.193 0.847 0.358
0.067 0.067 0.217
;
PROC PRINCOMP DATA=ACP OUT=pcstuff N=4 standard;
VAR ps psKha rend N P K Mg Ca Na AZ Fe B Cu Mn Zn;
RUN;
PROC CORR DATA=pcstuff;
WITH prin1 prin2 ;
RUN;
PROC FACTOR DATA=stuff SCREE;
RUN;
proc print;
VAR prin1 prin2 ;
run;
Tabla 10-2. Algoritmo para las variables fisiológicas de crecimiento (biomasa seca, área
foliar, consumo de nutrientes, uso eficiente de nutrientes) diseño en medidas repetidas
factorial incompleto en parcelas subdivididas-modelo mixto. Los factores entre sujetos
son anidados, pues la naturaleza de la matriz es incompleta.
DATA DMR;
INPUT LOCALIDAD $ CULTIVAR $ FERT $ REP p75 p100 p125 p150 @@;
DATALINES;
tubérculo
SUBACHOQUE CAPIRO CON 1 -1.13785 -0.46483 0.87791

0.98790
SUBACHOQUE CAPIRO SIN 1 -1.41514 -1.02719 -0.60019 -
0.37204
SUBACHOQUE SUPREMA CON 1 -1.2605 -0.30748 -0.10947 1.65284
SUBACHOQUE SUPREMA SIN 1 -1.23481 -0.66493 -0.17358
0.41534
FACATATIVA CAPIRO CON 1 -0.99081 0.05344 0.45485
1.29750
FACATATIVA CAPIRO SIN 1 -1.15979 -0.62750 0.35562
0.95927
FACATATIVA SUPREMA CON 1 -1.34376 -0.95720 -0.04522
0.53148
FACATATIVA SUPREMA SIN 1 -1.44174 -0.92939 -0.06431
0.65126
0.97930
SUBACHOQUE CAPIRO SIN 2 -1.47789 -0.93219 -0.58131 -
0.10330
SUBACHOQUE SUPREMA CON 2 -1.09168 -0.35672 -0.03050
1.92038
0.67275
FACATATIVA CAPIRO CON 2 -1.05123 -0.26630 0.46260
2.03499
1.06888
0.21719
0.32781
1.08567
SUBACHOQUE CAPIRO SIN 3 -1.41937 -0.99279 -0.54504
0.00373
SUBACHOQUE SUPREMA CON 3 -1.04724 -0.20945 -0.02044
1.84423
0.70618
FACATATIVA CAPIRO CON 3 -1.15326 -0.16141 0.17493
1.91770
1.05839
0.66164
0.23400
CHOCONTA CAPIRO CON 1 -0.96543 0.14505 0.72708
2.51486
CHOCONTA CAPIRO SIN 1 -0.60440 -0.22267 0.15680
0.91650
boyacense
CHOCONTA SUPREMA CON 1 -0.41776 0.48075 1.08363

2.57793
CHOCONTA SUPREMA SIN 1 -0.45914 0.18625 0.69411
2.03994
CHOCONTA CAPIRO CON 2 -0.60368 -0.20507 0.33177
1.95313
CHOCONTA CAPIRO SIN 2 -0.60440 -0.22267 0.15680
0.91650
CHOCONTA SUPREMA CON 2 -0.45483 0.12484 0.64641 2.96014
CHOCONTA SUPREMA SIN 2 -0.41776 0.48075 1.08363 2.57793
CHOCONTA CAPIRO CON 3 -0.99376 -0.34586 0.60951
1.90445
CHOCONTA CAPIRO SIN 3 -0.76955 -0.47609 -0.14430
0.12684
CHOCONTA SUPREMA CON 3 -0.52010 0.71076 1.21645
1.93838
CHOCONTA SUPREMA SIN 3 -0.71832 0.02954 0.70800
1.53854
;
*PROC PRINT;
*RUN;
title2 'DISEÑO EN MEDIDAS REPETIDAS(FACTORIAL INCOMPLETO EN PARCELAS
SUBDIVIDIDAS)';
proc glm data = DMR;
class LOCALIDAD CULTIVAR FERT REP ;
model p75 p100 p125 p150 = REP LOCALIDAD CULTIVAR LOCALIDAD*CULTIVAR
FERT(LOCALIDAD*CULTIVAR) / nouni ;
repeated time 4 profile/printe ;
TEST H=LOCALIDAD E=REP*LOCALIDAD;
TEST H=CULTIVAR E=REP*LOCALIDAD*CULTIVAR;
TEST H=FERT E=REP*LOCALIDAD*CULTIVAR;
run;

incompleto en parcelas subdivididas-modelo mixto inter sujetos para los componentes
que reunía la variables de mayor significancia .
Cuadrado de la
Fuente DF Tipo III SS F-Valor Pr > F
media
REP 2 0.11874366 0.05937183 1.00 0.3827
LOCALIDAD 2 1.579.091.282 789.545.641 133.48 <.0001
CULTIVAR 1 0.17788501 0.17788501 3.01 0.0969
LOCALIDAD*CULTIVAR 2 781.140.666 390.570.333 66.03 <.0001
FERT(LOCALI*CULTIVA) 6 866.295.486 144.382.581 24.41 <.0001
Error 22 130.132.949 0.05915134
tubérculo

incompleto en parcelas subdivididas-modelo mixto intrasujetos para los componentes que
reunía la variables de mayor significancia.
Cuadrado de Adj Pr > F

Fuente DF Tipo III SS F-Valor Pr > F
la media G-G H-F
Fenología 3 9.781.097.008 3.260.365.669 1006.22 <.0001 <.0001 <.0001
Fenología*Rep 6 0.21564404 0.03594067 1.11 0.3666 0.3615 0.3666
Fenol*Localid 6 101.281.480 0.16880247 5.21 0.0002 0.0032 0.0003
Fenología*Cult 3 0.36505300 0.12168433 3.76 0.0149 0.0399 0.0165
Fenol*Local*Cult 6 291.564.200 0.48594033 15.00 <.0001 <.0001 <.0001
Fenol*fert
18 521.395.502 0.28966417 8.94 <.0001 <.0001 <.0001
(Local*Cult)
Error(time) 66 213.852.948 0.03240196
Tabla 10-5. Matriz de correlación de Pearson de los componentes de alta significancia .
% PSt PSt PFt N P K Mg Ca Na S
% PSt 1 0.6519 0.4861 0.4490 0.3808 0.5767 0.5823 0.5810 0.6506 0.6167
PSt 0.6519 1 0.9648 0.9124 0.8259 0.9768 0.9653 0.8923 0.9994 0.9559
PFt 0.4861 0.9648 1 0.9360 0.8967 0.9787 0.9457 0.8693 0.9649 0.9266
N 0.4490 0.9124 0.9360 1 0.8538 0.9425 0.9271 0.8657 0.9126 0.8763
P 0.3808 0.8259 0.8967 0.8538 1 0.8936 0.7910 0.8589 0.8264 0.7501
K 0.5767 0.9768 0.9787 0.9425 0.8936 1 0.9675 0.8993 0.9775 0.9471
Mg 0.5823 0.9653 0.9457 0.9271 0.7910 0.9675 1 0.8464 0.9658 0.9812
Ca 0.5810 0.8923 0.8693 0.8657 0.8589 0.8993 0.8464 1 0.8933 0.8029
Na 0.6506 0.9994 0.9649 0.9126 0.8264 0.9775 0.9658 0.8933 1 0.9569
S 0.6167 0.9559 0.9266 0.8763 0.7501 0.9471 0.9812 0.8029 0.9569 1

boyacense
Tabla 10-6. Coeficiente b del modelo lineal de la extracción de nutrientes N, P y K (

kg/ha) respecto al rendimiento.
Intervalo de confianza
Cultivar Seb
b 95%
Capiro (0,00221;0,00248) 6,71362E-05

N
Suprema (0,00189;0,00233) 0,000110095
Capiro (0,00046;0,00056) 2,57036E-05
P
Suprema (0,00029;0,00036) 1,6226E-05
Capiro (0,00506;0,00563) 0,000143191
K
Suprema (0,00509;0,00557) 0,000120026

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Acumulación y distribución de

macronutrientes minerales en dos

Manuel Iván Gómez Sánchez

Universidad Nacional de Colombia

Manuel Iván Gómez Sánchez

Doctor en Ciencias Agrarias

Ph.D. Stanislav Magnitskiy

Ph.D Luis Ernesto Rodríguez

Universidad Nacional de Colombia

A mi esposa, ayuda idónea, hermosa sabia y

A mis padres, hermanos y mi padre espiritual

“Como el Padre me ha amado, así también

Jesus de Nazaret tu Salvador Juan 15:9

Agradezco a la Facultad de Ciencias Agrarias de la Unal-Bogotá y el soporte dado en

Agradezco a la empresa privada Ingeplant Ingenieria en Nutrición de Cultivos SAS

Agradezco y exalto especialmente el soporte técnico y apoyo logístico en cultivo del

Palabras clave: uso eficiente de nutrientes, relaciones alométricas, consumo de lujo,

Key words: efficient use of nutrients, allometric relationships, luxury consumption,

Lista de figuras .............................................................................................................. XIV

2.5.1 Statistical analysis ..........................................................................................45

5. Diagnóstico de K+ y NO3ˉ en savia para determinar el estado nutricional en papa

Suprema in Typic Hapludands, Subachoque, Andic Eutrudepts, Facatativá, and Humic

Lista de Símbolos y abreviaturas

a Concentración del nutriente cuando el total de la biomasa es ≤ 1 t ha -1

Capiro Cultivar Diacol Capiro

CDC Curvas de dilución crítica

cv. (s) Cultivar (es)

DAF Duración del área foliar

dds Días después de siembra

dOFR Diferencia de la dosis óptima de fertilización

DWT Dry weight of tubers/Peso seco de tubérculos

EFN Eficiencia fisiológica del nutriente

Accumulated nutrients per tuber yield/Eficiencia de nutrientes para la

EtK Eficiencia de traslocación de potasio en el tubérculo

EtN Eficiencia de traslocación de nitrógeno en el tubérculo

EtP Eficiencia de traslocación de fósforo en el tubérculo

EUN Eficiencia de utilización del nutriente

FWT Fresh weight of tubers/Peso fresco de tubérculos

HEI harvest extraction index

IAF Índice del área foliar

IC/HI Índice de cosecha/Harvest index

IEC/EI Índice de extranción/Rate of nutrient extraction

INN Índice de nutrición

ISE Método de electrodo selectivo de iones

Kc Curva de dilución de potasio

Nc Curva de dilución de nitrógeno

Nutrient use efficiency in tubers/ Eficiencia de uso de nutrientes en

PFt Rendimeinto potencial

Pc Curva de dilución de fósforo

OFR Dosis óptima de fertilización

Recuperación de nutrientes del fertilizante por el tubérculo/Rates of

Subsp., spp Subespecie

Suprema Cultivar Pastusa Suprema

W Biomasa seca total

Para lograr lo anterior, en el ámbito nacional es necesario establecer estrategias que

Para el año 2016, en Colombia se logró una producción estimada de 2,623,700 t ha -1 en

Desde el punto de vista de la productividad agrícola, la nutrición mineral es uno de los

La fisiología de la nutrición mineral permite entender el funcionamiento de la planta y

El N es el elemento esencial para la síntesis de proteínas, respiración, crecimiento de

Estudios realizados por Villamil (2005) con respecto al comportamiento de la

(2005) encontraron para el cv. Alpha que la absorción de P es mayor al inicio de

El potasio (K) es esencial para la translocación de azúcares hacia los tubérculos,

La partición de nutrientes minerales en hojas, tallos y tubérculos durante el crecimiento

Capiro 9.02 W –0.269 1-22 0.91 7.78-10.3 0.19-0.34 0.6205 0.0400

K Suprema 6.585 W –0.1353 1-24 0.92 5.38-7.76 0.05-0.18 0.600 0.0399