© The Authors, 2025, Published by the Universidad del Zulia*Corresponding author: dmarmolejo@uncp.edu.pe
Keywords:
GCA
SCA
Diallel crosses
Heritability
Yield
Genetic variance
General, specic combining ability, and heritability in potato genotypes (Solanum tuberosum
L.) for agronomic traits
Aptitud combinatoria general, especíca y heredabilidad en genotipos de papa (Solanum tuberosum
L.) para características agronómicas
Capacidade geral, especíca de combinação e herdabilidade em genótipos de batata (Solanum
tuberosum L.) para características agronômicas
Doris Marmolejo-Gutarra*
Jhosellin Alghira Arias Requena
Mónica Patricia Marín Huarcaya
Rev. Fac. Agron. (LUZ). 2025, 42(4): e254249
ISSN 2477-9407
DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v42.n4.VI
Crop production
Associate editor: Dr. Jorge Vilchez-Perozo
University of Zulia, Faculty of Agronomy
Bolivarian Republic of Venezuela
Universidad Nacional del Centro del Perú, Facultad de
Agronomía. Mariscal Castilla 3909, Huancayo. Código postal
12006, Perú.
Received: 12-07-2025
Accepted: 29-09-2025
Published: 16-10-2025
Abstract
Potato (Solanum tuberosum L.) is a key crop in Peru due
to its nutritional value and its potential for agronomic genetic
improvement, which justies further research in the selection of
promising parents and crosses. This study aimed to estimate general
combining ability (GCA), specic combining ability (SCA), and
heritability for key agronomic traits: plant height, earliness, and
the number and weight of tubers per plant, under the climatic
conditions of Huancayo, Peru. Sixteen F₁ families derived from a
full diallel cross (Gring’s model I, method I, xed eects) among
four parents (Mariva, Redondo, Redondo Achatado, and Oblongo)
were evaluated. The trial was conducted using a randomized
complete block design with three replications. The analysis
included ANOVA, estimates of GCA, SCA, and reciprocal eects.
Highly signicant dierences (p < 0.01) were detected among the
combinations. Mariva showed the highest GCA for plant height
(5.266), while the Redondo Achatado × Redondo cross exhibited the
highest SCA (6.404); for earliness, the GCA of Redondo Achatado
(0.056) and the SCA of Redondo Achatado × Mariva (0.952) were
outstanding; regarding tuber number per plant, Redondo had the
highest GCA (3.258) and the Redondo × Redondo Achatado cross
the highest SCA (8.982); for tuber weight, Redondo had the best
GCA (0.107) and Oblongo × Redondo the highest SCA (0.449).
Heritability ranged from 0.419 to 0.596, indicating moderate to
high genetic variation. The high values of GCA and SCA, along
with the observed heritability, conrm the potential of these parents
to improve potato yield.
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Rev. Fac. Agron. (LUZ). 2025, 42(4): e254249 October-December. ISSN 2477-9409.
2-6 |
Resumen
La papa (Solanum tuberosum L.) es clave en el Perú por su valor
alimentario y su potencial de mejoramiento genético agronómico,
lo cual justica profundizar en la selección de progenitores y cruzas
promisorias. Este estudio tuvo como objetivo estimar la aptitud
combinatoria general (ACG), especíca (ACE) y la heredabilidad
para características agronómicas clave: altura de planta, precocidad,
número y peso de tubérculos por planta, bajo condiciones climáticas
de Huancayo, Perú. Se evaluaron dieciséis familias F₁ obtenidas
mediante un cruzamiento dialélico completo (método I, modelo I de
Gring, efectos jos) entre cuatro progenitores (Mariva, Redondo,
Redondo Achatado y Oblongo). El experimento se condujo en un
diseño de bloques completos al azar con tres repeticiones. El análisis
incluyó ANDEVA, estimaciones de ACG, ACE y efectos recíprocos.
Se detectaron diferencias altamente signicativas (p < 0,01) entre
combinaciones. Mariva mostró la mayor ACG en altura de planta
(5,266) y la cruza Redondo Achatado × Redondo la máxima ACE
(6,404); en precocidad destacaron la ACG de Redondo Achatado
(0,056) y la ACE de Redondo Achatado × Mariva (0,952); en número
de tubérculos por planta sobresalieron la ACG de Redondo (3,258)
y la ACE de Redondo × Redondo Achatado (8,982); para peso de
tubérculos resaltaron la ACG de Redondo (0,107) y la ACE de
Oblongo × Redondo (0,449). La heredabilidad osciló entre 0,419 y
0,596, indicando variación genética de magnitud intermedia-alta.
Los valores elevados de ACG y ACE, junto con la heredabilidad
observada, conrman el potencial de estos progenitores para mejorar
el rendimiento de la papa.
Palabras clave: ACG, ACE, cruzas dialélicas, heredabilidad,
rendimiento, varianza genética.
Resumo
A batata (Solanum tuberosum L.) é um cultivo fundamental
no Peru devido ao seu valor alimentar e ao seu potencial para o
melhoramento genético agronômico, o que justica aprofundar a
seleção de genitores e cruzamentos promissores. Este estudo teve
como objetivo estimar a capacidade geral de combinação (CGC),
capacidade especíca de combinação (CEC) e a herdabilidade para
características agronômicas-chave: altura da planta, precocidade,
número e peso de tubérculos por planta, sob as condições climáticas
de Huancayo, Peru. Foram avaliadas dezesseis famílias F₁ obtidas
por meio de um cruzamento dialélico completo (Método I, Modelo
I de Gring, efeitos xos) entre quatro genitores (Mariva, Redondo,
Redondo Achatado e Oblongo). O experimento foi conduzido em
delineamento de blocos ao acaso com três repetições. A análise
incluiu ANOVA, estimativas de CGC, CEC e efeitos recíprocos.
Detectaram-se diferenças altamente signicativas (p < 0,01) entre
as combinações. Mariva apresentou a maior CGC para altura da
planta (5,266) e o cruzamento Redondo Achatado × Redondo obteve
a maior CEC (6,404); para precocidade, destacaram-se a CGC de
Redondo Achatado (0,056) e a CEC de Redondo Achatado × Mariva
(0,952); para número de tubérculos por planta, a CGC de Redondo
(3,258) e a CEC de Redondo × Redondo Achatado (8,982) foram
superiores; para peso de tubérculos, sobressaíram a CGC de Redondo
(0,107) e a CEC de Oblongo × Redondo (0,449). A herdabilidade
variou entre 0,419 e 0,596, indicando variação genética de magnitude
intermediária a alta. Os altos valores de CGC e CEC, juntamente com
a herdabilidade observada, conrmam o potencial desses genitores
para o aumento do rendimento da batata.
Palavras-chave: CGC, CEC, cruzamentos dialélicos, herdabilidade,
rendimento, variância genética.
Introduction
Potato (Solanum tuberosum L.) is a strategic crop in Peru, both for
its nutritional value and for its economic relevance. Peru, the Andean
region, is one of the main centers of origin and diversication of this
species; it is home to a great genetic wealth of native and improved
cultivars (Ovchinnikova et al., 2011). In 2023, Peru recorded a
production of more than 5.1 million tons, despite having faced rainfall
deciency that year (MIDAGRI, 2024), positioning itself as the largest
potato producer in Latin America. However, its average yield per
hectare is still low regarding its genetic potential, due to factors such
as limited genetic variability in cultivated materials, susceptibility to
biotic and abiotic stresses, and poor adaptation to changing conditions
(MIDAGRI, 2018, 2024). In this context, genetic improvement is a
priority tool to increase crop productivity, adaptability, and quality,
which justies the need to identify superior parents and promising
crosses through genetic analyses such as general combining ability
(GCA), specic combining ability (SCA), and heritability.
The diallel design, proposed by Gring (1956), allows the
simultaneous estimation of general combining ability (GCA) and
specic combining ability (SCA), providing essential information on
the additive and non-additive eects that control agronomic traits of
interest. GCA is mainly associated with the additive action of genes,
which makes it useful for the recurrent selection of superior parents
(Saavedra et al., 2021), while SCA reects the specic interaction
between parental combinations, making it possible to identify hybrids
with unexpected superior behavior due to dominance eects or
epistasis (Mugisa et al., 2022).
Several studies in potatoes and other crops indicate that, for
yield-related traits such as the number and weight of tubers per plant,
the non-additive eects may exceed additive eects in magnitude
(Amiri et al., 2020; Mugisa et al., 2022), especially in heterogeneous
environments. However, the heritability of the trait directly inuences
the eectiveness of selection: when it is high, prioritizing GCA is
recommended, whereas under conditions of moderate or low
heritability, SCA can oer advantages for direct cloning (Onofri et
al., 2021; Russell & Sandall, 2005).
In the case of potato, a tetraploid and allogamous species, the
genetic complexity is high, which justies the use of models that
consider xed genetic components. Gring’s model I, method
I, is appropriate for this type of analysis, as it also allows for the
estimation of reciprocal eects and components of genetic variance
useful for calculating narrow-sense heritability (Pooni et al., 1984;
Mohammed et al., 2016).
Despite advances in breeding programs, information on specic
hybrid combinations with agronomic potential under high Andean
conditions remains limited. Therefore, the present study aimed to
estimate GCA, SCA, reciprocal eects, and heritability in four potato
genotypes for agronomic traits such as plant height, earliness, and
number and weight of tubers per plant to identify promising parents
and crosses for inclusion in genetic improvement programs.
Materials and methods
The research was carried out during the 2023-2024 agricultural
season in two phases: the generation of F₁ hybrids in greenhouses
and the eld evaluation of the resulting progeny. The controlled
crosses were performed in the greenhouse of the Agricultural
Sciences Research Center (CICA) of Huancayo, while the eld phase
was carried out in the district of Quichuay, province of Huancayo,
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Marmolejo-Gutarra et al. Rev. Fac. Agron. (LUZ). 2025, 42(4): e254249
3-6 |
The eld experiment was conducted under a randomized complete
block design (RCBD) with 16 families and three replications. All
possible combinations of the complete diallel cross among the four
parents were evaluated (method I, Gring’s model I with xed genetic
eects): 4 self-fertilizations, 6 direct crosses, and 6 reciprocal crosses
(Gring, 1956). Each treatment (hybrid family and self-fertilized
parent) was established in one plot per block, with 15 plants per plot
(45 plants per family in total). A total of 720 plants were transplanted
in an experimental area of 216 m². During the crop cycle, standard
cultural practices for potatoes were carried out, including hilling,
manual weed control on two occasions, and preventive applications
against insects and pathogens.
Variables evaluated
Three important agronomic variables were measured: plant
height at 120 days after transplanting, number of tubers per plant, and
total tuber weight per plant in kilograms. These measurements were
averaged at the plot and treatment level.
Statistical analysis
The collected data were initially subjected to an analysis of
variance (ANDEVA) under the randomized complete block design
model. Subsequently, the diallel analysis was carried out according
to Gring’s model I, method I (1956), considering parents, direct
and reciprocal crosses. This analysis allowed for the estimation of
general combining ability (GCA), specic combining ability (SCA),
and reciprocal eects.
The general statistical model applied was:
Y₍ᵢⱼₖ₎ = μ + gᵢ + sᵢⱼ + mᵢ + lᵢⱼ + Bₖ + eᵢⱼₖ
Where: Y₍ᵢⱼₖ₎ is the observed value; μ, the overall mean; g
i
, the
eect of GCA of parent i; s
ij
, the eect of SCA between parents i and
j; e, the maternal eect; l
ij
, the reciprocal eect; B
k
, the block eect;
and e
ijk
, the random error.
Estimated eects:
ACGᵢ = Ȳᵢ. − μ
ACEᵢⱼ = Ȳᵢⱼ − μ − ACGᵢ − ACGⱼ
Variances and narrow-sense heritability (h²):
Additive variance:
Dominance variance:
Total phenotypic variance:
Heritability (h²):
All statistical analyses were carried out using the InfoGen version
2018 software (Balzarini & Di Rienzo, 2003) and Microsoft Excel
2019 for organization and preliminary data processing.
department of Junín, Peru. The experimental eld is located at 3.606
m.a.s.l., with a clay-loam soil texture (pH 7.1) and an Andean climate
characterized by temperatures ranging between 4 and 18 °C.
The genetic material was made up of four potato genotypes:
Redondo, Redondo Achatado, Oblongo (Solanum tuberosum L.),
and Mariva (Solanum tuberosum subsp. andigenum). These parents,
provided by CICA, were established from seed tubers. Male parents
were established in pots to induce owering and collect pollen, while
female parents were managed using the brick method (gure 1). This
method consisted of making cuts in the stolons during the owering
phase to prolong it, facilitate manual pollination, and improve the
production of botanical seeds.
Figure 1. Planting tubers using the brick method.
Once the parent plants entered owering, controlled crosses were
made by hand pollination. Prior to pollination, manual emasculation of
the bud owers of the female parents was performed, extracting their
stamens before dehiscence to prevent self-fertilization. Immediately
after, fresh pollen collected from the male parents was applied to
the stigmas of the emasculated owers. Each pollinated ower was
labeled with the identication of the cross (female parent × male
parent), and the fruit (berry) was allowed to develop on the mother
plant. Approximately 6-8 weeks after pollination, when the berries
reached maturity (darkening of the epidermis), they were harvested
manually. From the harvested berries, the sexual seeds corresponding
to each cross were extracted (table 1), which were washed and dried
at room temperature for temporary storage.
Table 1. Codes of hybrid potato clones.
Treatment
N°
Parental
genotype used
Crossbreeding of origin Clonal code
1 Mariva Mariva × Vacapa Ccallum IPC 720025
2 Redondo Yungay × Mariva GOP-031513.02
3
Redondo
Achatado
Yungay × Mariva GOP-031513.01
4 Oblongo Yungay × Perricholi GOP-031512.09
IPC: International Potato Center, “720025” corresponds to the access code of the IPC
genebank/material; Clonal code (example: GOP-031513.02): GOP = plant breeder initials,
03 = breeding code, 15 = year of crossbreeding, 1 = female parent, 3 = male parent, and 02 =
breeding number within the progeny.
The botanical seeds from each cross were sown in seedbeds
(trays lined with paper towels). Once germinated, the seedlings were
transplanted into vessels with substrate and kept in a nursery until
they reached around 10 cm in height (with 2-3 true leaves). Next, the
seedlings were transplanted in the nal eld of Quichuay, in rows 0.90 m
apart, with a distance of 0.30 m between plants (gure 2). Immediately
after transplanting, establishment irrigation was carried out, and bottom
fertilizers were applied to ensure a good start of the crop.
Figure 2. a: Transplanting of seedlings to the nal eld. b:
Harvesting of hybrid potato tubers in the second
cultivation phase.
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Rev. Fac. Agron. (LUZ). 2025, 42(4): e254249 October-December. ISSN 2477-9409.
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Table 4. General and specic combining ability eects for the
number of tubers per plant evaluated at harvest (120
days after planting).
Parents
Mariva
(1)
Redondo
(2)
Redondo
Achatado (3)
Oblongo
(4)
Mariva (1) -0.296 -4.151 -6.281 -2.864
Redondo (2) 7.052* 3.258 8.982** 4.560
Redondo Achatado (3) -1.326 1.679 -4.092 7.347*
Oblongo (4) -0.967 -0.673 2.160 1.130
* = signicant at 5 %; ** = signicant at 1 %. Values on the main diagonal correspond to the
eects of the general combining ability GCA of each parent. O-diagonal values represent
the eects of the specic combining ability SCA of each hybrid combination. Positive values
indicate favorable eects on the evaluated variable, and negative values, unfavorable eects.
Tuber weight per plant
Regarding the weight of tubers per plant (table 5), Redondo showed
the best GCA (0.107), highlighting its additive genetic contribution to
the development of tuber weight. The hybrid combinations Oblongo ×
Redondo (0.449), Mariva × Redondo Achatado (0.403), and Mariva ×
Oblongo (0.101) stood out in SCA, revealing important non-additive
eects possibly linked to epistatic or dominant interactions. These
ndings are in agreement with what was reported by Mohammed et.
al. (2016), who point out that the yield per plant in potatoes depends
signicantly on non-additive eects.
Table 5. General and specic combining ability eects for tuber
weight per plant evaluated at harvest (120 days after
planting).
Parents
Mariva
(1)
Redondo
(2)
Redondo
Achatado (3)
Oblongo
(4)
Mariva (1) -0.059 -0.119 0.217 -0.234
Redondo (2) 0.079 0.107 0.007 -0.226
Redondo Achatado (3) 0.403 0.014 -0.110 0.013
Oblongo (4) 0.101 0.449* -0.287 0.063
* = signicant at 5%; ** = signicant at 1%. Values on the main diagonal correspond to the
eects of the general combining ability GCA of each parent. O-diagonal values represent
the eects of the specic combining ability SCA of each hybrid combination. Positive values
indicate favorable eects on the evaluated variable, and negative values, unfavorable eects.
Tukey’s mean test for evaluated agronomic variables
In table 6, the results of Tukey’s test (p < 0.05) reveal statistically
signicant dierences between the 16 hybrid families evaluated,
which shows considerable genetic variability for the agronomic traits
studied.
For plant height, the combination Mariva × Mariva (84.31 cm)
was statistically superior to the rest of the families, followed by
Oblongo × Redondo (82.81 cm), and Redondo × Mariva (82.63 cm).
These results suggest a possible favorable additive eect on height
transmitted by the parent Mariva, as well as specic combinations
with positive non-additive eects, especially when Oblongo is used
as a female parent. Darabad et al. (2020) point out that a higher plant
height could be associated with greater vigor and eciency in light
capture, favoring foliar and photosynthetic development, a desirable
condition in high-altitude environments such as Huancayo (3200
m.a.s.l.).
In relation to earliness, it was identied that the crosses Redondo
Achatado × Mariva (2.82) and Mariva × Redondo Achatado (2.81)
presented a greater number of days until senescence, indicating a
longer duration of the vegetative cycle. This behavior could reect
a cumulative additive gene action in favor of delayed maturity,
which may be advantageous in areas where the agricultural cycle
Results and discussion
Diallel analysis allowed the decomposition of observed variance
into additive genetic eects (GCA), non-additive eects (SCA), and
reciprocal eects (table 2). Highly signicant eects (p<0.01) of
GCA and SCA were detected in the evaluated variables, indicating
the joint participation of additive and non-additive gene action in the
expression of these traits. Signicant reciprocal eects (p<0.05) were
also observed in plant height, number, and weight of tubers per plant,
suggesting the inuence of maternal or cytoplasmic eects on these
variables.
Table 2. Analysis of variance (ANAVA) of Gring’s method for
tubers in potato (Solanum tuberosum L.) genotypes and
hybrids.
Source of
variation
Gl
Plant height Earliness
Number of
tubers
Weight of
tubers
(CM) (CM) (CM) (CM)
GCA 3 141.094** 0.042ns 76.592** 0.083*
SCA 6 27791.964** 0.677** 20919.638** 0.33**
Reciprocal 6 44.764** 0.045ns 73.443** 0.056*
Error 30 12.191 0.024 15.712 0.022
Plant height
In plant height (table 3), Mariva stood out with a high general
combining ability (GCA) of 5.266, indicating a strong ability to
transmit genes related to vegetative vigor, in accordance with previous
studies in potatoes carried out by Khan et al. (2013). Likewise, the
crosses Redondo × Redondo Achatado (6.404), Redondo × Oblongo
(4.870), and Mariva × Redondo Achatado (3.128) showed signicant
values of specic combining ability (SCA), indicating favorable
non-additive interactions for this trait. As indicated by Luthra et al.
(2005), additive eects are essential for vegetative traits, but specic
combinations may provide additional advantages.
Table 3. General and specic combining ability eects for plant
height evaluated at harvest (120 days after planting).
Parents Mariva (1) Redondo (2)
Redondo
Achatado (3)
Oblongo
(4)
Mariva (1) 5.266* -5.593 4.205 -4.276
Redondo (2) -1.396 1.151 3.018 -6.697
Redondo
Achatado (3)
3.128 6.404* -4.487 -3.617
Oblongo (4) -3.497 4.870 -0.034 -1.930
* = signicant at 5 %; ** = signicant at 1 %. Values on the main diagonal correspond to the
eects of the general combining ability GCA of each parent. O-diagonal values represent
the eects of the specic combining ability SCA of each hybrid combination. Positive values
indicate favorable eects on the evaluated variable, and negative values, unfavorable eects.
Number of tubers per plant
For the number of tubers (table 4), the Redondo parent had the
highest GCA (3.258), reecting its ability to transmit prolicacy.
In SCA, the combinations Redondo Achatado × Redondo (8.982),
Oblongo × Redondo Achatado (7.347), and Mariva × Redondo
(7.052) stood out. These results are consistent with Onofri et al.
(2021), who suggest that combining parents with high GCA and SCA
optimizes selection for yield-related traits. This is also consistent with
specic ndings in potato by Kamara et. al. (2021), who recommend
dual selection strategies.
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Marmolejo-Gutarra et al. Rev. Fac. Agron. (LUZ). 2025, 42(4): e254249
5-6 |
is extended. In contrast, Mariva × Mariva, Oblongo × Mariva, and
Oblongo × Redondo Achatado showed the lowest values (<1.25),
evidencing a genetic pattern oriented towards early maturity. This is
consistent with what was reported by Khan et al. (2013), who state
that additive gene action has a greater inuence on the determination
of physiological maturity in potatoes.
Regarding the number of tubers per plant, the cross Redondo ×
Mariva reached the highest average (76.64), followed by Redondo
× Redondo Achatado (72.30) and Redondo × Oblongo (70.74). This
pattern suggests a positive eect of the Redondo parent when used as
a male, reecting its favorable specic combining ability. According
to Onofri et al. (2021), the SCA is key to identifying combinations of
parents that exceed the expected yield of their individual GCA values,
which is observed in these crosses. In addition, the result aligns with
the ndings of Mugisa et al. (2022), who observed in sweet potato that
certain hybrid combinations with high SCA values were candidates
for direct cloning in vegetative propagation programs.
Finally, for the weight of tubers per plant, the family Oblongo ×
Redondo showed the highest value (2.42 kg), followed by Mariva
× Redondo Achatado (2.03 kg), and Oblongo × Mariva (1.92 kg).
These combinations reect not only favorable specic eects but also
the importance of the maternal eect on total yield. As highlighted
by Saavedra et al. (2021), reciprocal eects may be relevant when
the female parent inuences the phenotypic expression of complex
traits such as yield. Likewise, Mackay et al. (2021) state that
optimal biomass utilization is explained by the recombination and
redistribution of favorable alleles during sexual reproduction. Taken
together, these results reinforce the importance of considering both
additive (GCA) and non-additive (SCA) and reciprocal eects when
selecting parents for potato breeding programs, especially under
high-altitude agroecological conditions such as in Huancayo.
Estimation of variance and heritability of height, earliness,
and yield per plant evaluated at harvest (120 days after planting)
Table 7 presents the estimates of additive variance, dominant
variance, and narrow-sense heritability (h²) for the traits of plant
height, earliness, number and weight of tubers per plant, based on the
diallel design proposed by Gring (1956).
Table 6. Signicance test of the means for plant height (cm), earliness, number of tubers per plant, and tuber weight (kg).
O.M. Families Plant height (cm) Earliness Tubers.plant
-1
Weight (kg) tubers.plant
-1
1 1×1 (M × M) 84.31 ± 1.56ᵃ 1.22 ± 0.08ᵈ 57.12 ± 1.33ᵃᵇᶜᵈ 0.88 ± 0.06ᵈ
9 1×2 (M × R) 71.45 ± 1.56ᵃᵇᶜᵈ 1.83 ± 0.08ᵇᶜᵈ 68.33 ± 1.33ᵃᵇᶜᵈ 1.59 ± 0.06ᵇᶜᵈ
4 1×3 (M × RA) 80.13 ± 1.56ᵃᵇᶜ 2.81 ± 0.08ᵃ 50.48 ± 1.33ᵈ 2.03 ± 0.06ᵃᵇ
12 1×4 (M × O) 67.58 ± 1.56ᵃᵇᶜᵈ 1.82 ± 0.08ᵇᶜᵈ 59.48 ± 1.33ᵃᵇᶜᵈ 1.45 ± 0.06ᵇᶜᵈ
3 2×1 (R × M) 82.63 ± 1.56ᵃᵇ 1.51 ± 0.08ᶜᵈ 76.64 ± 1.33ᵃ 1.83 ± 0.06ᵃᵇᶜ
14 2×2 (R × R) 64.44 ± 1.56ᵇᶜᵈ 1.51 ± 0.08ᶜᵈ 60.93 ± 1.33ᵃᵇᶜᵈ 1.25 ± 0.06ᵇᶜᵈ
5 2×3 (R × RA) 78.11 ± 1.56ᵃᵇᶜ 1.67 ± 0.08ᵇᶜᵈ 72.30 ± 1.33ᵃᵇ 1.60 ± 0.06ᵇᶜᵈ
10 2×4 (R × O) 69.41 ± 1.56ᵃᵇᶜᵈ 2.14 ± 0.08ᵃᵇᶜ 70.74 ± 1.33ᵃᵇᶜ 1.97 ± 0.06ᵃᵇᶜ
8 3×1 (RA × M) 71.72 ± 1.56ᵃᵇᶜᵈ 2.82 ± 0.08ᵃ 63.04 ± 1.33ᵃᵇᶜᵈ 1.60 ± 0.06ᵇᶜᵈ
7 3×2 (RA × R) 72.07 ± 1.56ᵃᵇᶜ 1.84 ± 0.08ᵇᶜᵈ 54.33 ± 1.33ᵇᶜᵈ 1.58 ± 0.06ᵇᶜᵈ
16 3×3 (RA × RA) 53.55 ± 1.56ᵈ 1.60 ± 0.08ᵇᶜᵈ 51.77 ± 1.33ᶜᵈ 1.23 ± 0.06ᶜᵈ
15 3×4 (RA × O) 61.95 ± 1.56ᶜᵈ 1.11 ± 0.08ᵈ 69.02 ± 1.33ᵃᵇᶜᵈ 1.26 ± 0.06ᵇᶜᵈ
6 4×1 (O × m) 76.13 ± 1.56ᵃᵇᶜ 1.22 ± 0.08ᵈ 65.20 ± 1.33ᵃᵇᶜᵈ 1.92 ± 0.06ᵃᵇᶜ
2 4×2 (O × R) 82.81 ± 1.56ᵃᵇ 2.35 ± 0.08ᵃᵇ 70.74 ± 1.33ᵃᵇᶜ 2.42 ± 0.06ᵃ
11 4×3 (O × RA) 69.19 ± 1.56ᵃᵇᶜᵈ 1.22 ± 0.08ᵈ 54.32 ± 1.33ᵇᶜᵈ 1.23 ± 0.06ᶜᵈ
13 4×4 (O × O) 66.82 ± 1.56ᵃᵇᶜᵈ 1.75 ± 0.08ᵇᶜᵈ 64.21 ± 1.33ᵃᵇᶜᵈ 1.44 ± 0.06ᵇᶜᵈ
Dierent letters in each column indicate signicant dierences according to Tukey’s test (p < 0.05). Redondo (R), Redondo Achatado (RA), Oblongo (O), Mariva (M).
The results show that, in all the traits evaluated, dominance
eects predominate, observing that the dominant variance (σ²D)
was consistently higher than the additive variance (σ²A). This trend
evidences the existence of non-additive eects such as dominance and
epistasis in the genetic expression of the evaluated traits, coinciding
with what was reported by Li et al. (2010), who pointed out that many
traits in potato yield respond to complex gene interactions. In the case
of plant height, a narrow-sense heritability of 0.5951 was obtained,
which indicates a moderate to high capacity to transmit this trait to the
progeny. This suggests that this trait may respond eciently to early
selection in breeding programs. Regarding earliness, the h² value was
0.5718, which also represents a considerable genetic potential, being
favorable for selecting early lines adapted to short production cycles
or limiting climatic conditions. According to Schmidt et al. (2019),
this level of heritability allows for eective genetic advancement
when combined with adequate environmental control.
For the number of tubers per plant, a heritability of 0.5969 was
recorded, which denotes an important genetic inuence. This is
consistent with what was reported by Maibvisira et al. (2018), who
found high heritability for yield components, particularly in crosses
with well-contrasted parents. In contrast, the weight of tubers per
plant had the lowest heritability (0.4190), which indicates that this
trait is more inuenced by the environment and by non-additive
genetic eects. This result suggests that, to improve weight, it is
necessary to employ strategies such as the selection of specic hybrid
combinations of high SCA, and to perform multifocal evaluations
to control the environmental eect. This approach is supported by
Mondal et al. (2022), who state that when environmental variance
is low compared to genetic variance, genetic control can be more
reliably exploited. The estimated values support the use of mixed
breeding strategies, combining phenotypic selection based on GCA
with the identication of specic crosses with high SCA. This allows
for the utilization of both moderate heritability and dominance eects,
which are key to developing more productive clonal cultivars better
adapted to the agroecological conditions of the Mantaro Valley.
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Rev. Fac. Agron. (LUZ). 2025, 42(4): e254249 October-December. ISSN 2477-9409.
6-6 |
Table 7. Analysis of genetic components and heritability of
height, earliness, and yield.
Trait
Additive
Variance (σ²A)
Dominant
variance (σ²D)
Heritability
(h²)
Plant height 6378.49100 17095.24500 0.59510
Earliness 0.14600 0.40200 0.57180
N° of tubers/plant 4808.76300 12863.95500 0.59690
Tuber/plant weight 0.05600 0.18900 0.41900
Conclusions
The genetic combinations Redondo × Redondo Achatado,
Oblongo × Redondo, and Redondo × Oblongo showed highly
signicant SCA eects that increased plant height, earliness, number,
and weight of tubers per plant.
Narrow-sense heritability was moderate to high for height
(0.595), number of tubers (0.597), and earliness (0.572), conrming
an additive component that can be exploited by selecting parents with
high GCA.
In tuber weight (h² = 0.419), dominance eects predominated;
therefore, it is recommended to exploit heterosis through specic
crosses.
Overall, these results support a mixed scheme: selecting Redondo
and Oblongo as base parents and establishing the hybrids Redondo ×
Redondo Achatado and Oblongo × Redondo as commercial clones,
with the aim of increasing potato yields in the Mantaro Valley.
Literature cited
Amiri, S., Dehdar Masjedlou, B., Panahi, B., & Mohammadi, R. (2020).
Combining ability analysis of tuber yield and related traits in potatoes.
Genetika, 52. https://doi.org/10.2298/GENSR2001215D
Balzarini, M., & Di Rienzo, J. (2003). Info-Gen: Software para análisis estad’istico
de datos genéticos. Facultad de Ciencia Agropecuarias. Universidad
Nacional de Córdoba. Argentina. https://www.info-gen.com.ar/
Darabad, G. R., Hassandokht, M. R., Hassanpanah, D., & Mousavi, A. (2020).
Diallel Cross in Potato Cultivars (Solanum tuberosum L.) and Evaluation
of Their Progenies Under Decit Water Stress. Acta Agrobotanica, 73(2).
https://doi.org/10.5586/aa.7325
Gring, B. (1956). Concept of General and Specic Combining Ability in
Relation to Diallel Crossing Systems. Australian Journal of Biological
Sciences, 9(4), 463–493. https://doi.org/10.1071/bi9560463
Khan, M. F., Tabassum, N., Latif, A., Khaliq, A., & Malik, M. (2013).
Morphological characterization of potato (Solanum tuberosum L.)
germplasm under rainfed environment. African Journal of Biotechnology,
12(21), 3214–3223. https://doi.org/10.5897/AJB11.4293
Li, L., Paulo, M.-J., van Eeuwijk, F., & Gebhardt, C. (2010). Statistical epistasis
between candidate gene alleles for complex tuber traits in an association
mapping population of tetraploid potato. Theoretical and Applied Genetics,
121(7), 1303–1310. https://doi.org/10.1007/s00122-010-1389-3
Luthra, S. K., Gopal, J., Pandey, S. K., & Singh, B. P. (2005). Genetic parameters
and characters associations in tuberosum potatoes. Potato Journal, 32(3–
4). https://epubs.icar.org.in/index.php/PotatoJ/article/view/33501
Mackay, I. J., Cockram, J., Howell, P., & Powell, W. (2021). Understanding the
classics: The unifying concepts of transgressive segregation, inbreeding
depression and heterosis and their central relevance for crop breeding.
Plant Biotechnology Journal, 19(1), 26–34. https://doi.org/10.1111/
pbi.13481
Maibvisira, N. A., Gasura, E., Maphosa, M., & Nyakurwa, C. S. (2018). Genetic
basis and the current breeding eorts for quality protein maize in
Southern Africa. African Crop Science Journal, 26(4), 529–541. https://
doi.org/10.4314/acsj.v26i4.7
MIDAGRI. (2018). Plan estratégico para promoción y difusión de variedades
de papa para procesamiento liberadas por CIP en Perú. International
Potato Center. https://cipotato.org/publications/plan-estrategico-para-
promocion-y-difusion-de-variedades-de-papa-para-procesamiento-
liberadas-por-cip-en-peru/
MIDAGRI. (2024). Siembras y Perspectivas de la Producción. https://www.gob.
pe/institucion/midagri/informes-publicaciones/4344772-observatorio-de-
siembras-y-perspectivas-de-produccion-2023
Mohammed, A. M., El-Kader, A., & El-Hegazy, M. H. (2016). Combining ability
and gene action for potato yield and its components under normal and
heat stress conditions. Middle East Journal of Agriculture Research, 5(4),
543-556. https://doi.org/10.3390/agronomy11081450
Mondal, S., Ortiz, R., & Crespo-Herrera, L. (2022). Quantitative Approaches to
Plant Breeding Concepts, Strategies and Practical Applications. https://
doi.org/10.3389/978-2-88976-878-3
Mugisa, I., Karungi, J., Musana, P., Odama, R., Alajo, A., Chelangat, D. M.,
Anyanga, M. O., Oloka, B. M., Gonçalves dos Santos, I., Talwana, H.,
Ochwo-Ssemakula, M., Edema, R., Gibson, P., Ssali, R., Campos, H.,
Olukolu, B. A., da Silva Pereira, G., Yencho, C., & Yada, B. (2022).
Combining ability and heritability analysis of sweetpotato weevil
resistance, root yield, and dry matter content in sweetpotato. Frontiers in
Plant Science, 13. https://doi.org/10.3389/fpls.2022.956936
Onofri, A., Terzaroli, N., & Russi, L. (2021). Linear models for diallel crosses:
A review with R functions. Theoretical and Applied Genetics, 134(2),
585–601. https://doi.org/10.1007/s00122-020-03716-8
Ovchinnikova, A., Krylova, E., Gavrilenko, T-, Smekalova, T., Zhuk, M., Knapp,
S., & Spooner, D. M. (2011). Taxonomy of cultivated potatoes (Solanum
section Petota: Solanaceae). Botanical Journal of the Linnean Society,
165(2), 107-155. https://doi.org/10.1111/j.1095-8339.2010.01107.x
Pooni, H. S., Jinks, J. L., & Singh, R. K. (1984). Methods of analysis and the
estimation of the genetic parameters from a diallel set of crosses. Heredity,
52(2), 243–253. https://doi.org/10.1038/hdy.1984.26
Kamara, M. M., Ibrahim, K. M., Mansour, E., Kheir, A. M. S-. Germoush, M. O.,
Abd El- Moneim, D., Motawei, M. I., Alhusays, A. Y., Farid, M.A.,rehan,
M. A., & Rehan, m. ( 2011). Combining Ability and Gene Action
Controlling Grain Yield and Its Related Traints in Bread Wheat under
Heat Stress and Normal Conditions. Agronomy, 11(8), 1450. https://doi.
org/10.3390/agronomy11081450.
Saavedra Guevara, C., Pérez López, D. de J., González Huerta, A., Franco
Martínez, J. R. P., Rubí Arriaga, M., Ramírez Dávila, J. F., Saavedra
Guevara, C., Pérez López, D. de J., González Huerta, A., Franco Martínez,
J. R. P., Rubí Arriaga, M., & Ramírez Dávila, J. F. (2021). Métodos de
Gring: Revisión sobre su importancia y aplicación en tomejoramiento
convencional. Revista mexicana de ciencias agrícolas, 12(7), 1275–1286.
https://doi.org/10.29312/remexca.v12i7.3040
Schmidt, P., Hartung, J., Bennewitz, J., & Piepho, H.-P. (2019). Heritability in
Plant Breeding on a Genotype-Dierence Basis. Genetics, 212(4), 991–
1008. https://doi.org/10.1534/genetics.119.302134
