Open Agriculture.
2017; 2: 561–570
Research Article
Manuel A. Gastelo Benavides*, Luis Diaz, Gabriela Burgos, Thomas Zum Felde,
Merideth Bonierbale
Heritability for Yield and Glycoalkaloid Content
in Potato Breeding under Warm Environments
https://doi.org/10.1515/opag-2017-0059
received November 21, 2016; accepted September 19, 2017
Abstract: High temperatures affect potato production in
the tropics, putting tuber yield and quality at risk and
leading to increased glycoalkaloid concentration the
cause of the bitter taste in potatoes and a cause for concern
for human health. The International Potato Center (CIP),
has developed new heat tolerant clones which are heat
tolerant and also resistant to late blight. These clones offer
an opportunity to evaluate yield and glycoalkaloid levels
after growth under high temperature environments. We
evaluated four sets of 16 full-sib families and 20 clones for
tuber yield and glycoalkaloid content in order to estimate
narrow-sense and broad-sense heritability respectively.
We used a randomized complete block design replicated in
three locations in Peru; San Ramon, La Molina and Majes
At harvest, the number and weight of marketable and non-
marketable tubers were recorded. We analyzed samples of
tubers from each clone for glycoalkaloid content using
spectrophotometry. Narrow-sense heritability for tuber
yield, tuber number and average tuber weight were 0.41,
0.50 and 0.83, respectively, indicating that further gains
in breeding for heat tolerance will be possible. Broad-
sense heritability for glycoalkaloid content was 0.63 and
correlation with tuber yield was weak, r=0.33 and R²=0.11
(P<0.01). High heritability and weak correlation will
allow us to select clones with high tuber yield and low
glycoalkaloid content, to serve as candidate varieties and
parents in breeding programs.
Keywords: heat tolerance, potato, tuber yield, heritability,
glycoalkaloid content
*Corresponding author: Manuel A. Gastelo Benavides, International
Potato Center (CIP), Peru, E-mail: m.gastelo@cgiar.org
Luis Diaz, Gabriela Burgos, Thomas Zum Felde, Merideth Bonierba-
le, International Potato Center (CIP), Peru
Open Access. © 2017 Manuel A. Gastelo Benavides et al., published by De Gruyter Open.
Attribution 4.0 Public License.
1 Introduction
Climate change is already affecting weather patterns in
traditional potato growing areas, where unpredictable
rain fall and higher temperatures, increase pressure by
pests and diseases and put tuber yield and quality at risk
(Hijmans 2003; Levy and Veilleux 2007). High temperature
is one of the major abiotic stresses on potato production in
many areas in the world and especially in the tropics (Birch
et al. 2012), and may lead to a drastic reduction of tuber
yield. Yield reduction however is not the only indicator
of potato susceptibility to high temperature, since the
occurrence of secondary tuberization and tuber defects of
a physiological nature (Rykaczewska 2015), should also
be taken into account. Examples of the latter include:
reduced photosynthesis and increased respiration;,
inhibition of tuber initiation and growth; acceleration of
haulm growth and appearance of tuber disorders such as
second growth, cracks, sprouting, deformations, reduced
tuber dry matter content and raised glycoalkaloid content
(Levy and Veilleux 2007). High temperatures particularly
at night during tuberization, decrease partitioning to
tubers favoring haulm growth (Ewing 1981). High levels
of glycoalkaloid give potatoes an unpleasant flavor and
could be toxic to humans (Storey and Davies 1992). The
maximum allowed level is 20mg/100g of fresh weight
(FW), but levels below 7mg/100gFW are preferred in new
potato varieties (Jacob van Dam 2002).
Average tuber weight has been suggested by Jefferies
and MacKerron (1987) as an important factor influencing
tuber yield, and Thompson et al. (1983) have suggested
using average tuber weight as an indirect selection
criterion. Shung-Lin Zheng et al. (2016) show that there
is negative correlation between tuber number and
harvestable yield in fall and spring seasons and the yield
components differed slightly in their contribution to
overall yield.
Luthra et al. (2013), estimated ten genetic parameters
in 105 potato genotypes during 2007 and 2008 under
high temperature stress conditions in an effort to aid
This work is licensed under the Creative Commons
Unauthenticated
Download Date | 11/19/17 1:06 AM
562   M.A. Gastelo Benavides, et al.
identification of superior parents with high tuber yield.
These authors found high genetic variability, heritability
and genetic advance for tuber yield and its components
while total tuber yield has a significant positive
correlation with marketable tuber yield and average
tuber weight. Tuber number were positively correlated
with stem number and negatively correlated with average
tuber weight. The results showed that parental lines could
be selected based on tuber yield and its components.
But other estimates of heritability under conditions of
unfavorable environments with high temperatures were
low; Gandolfi and Pereira (2011) recommended using
other selection criteria that are correlated with tuber
yield as indicators of heat tolerance.
Van Dam et al. (1999) found high heritability estimates
for glycoalkaloid content and no association with a large
number of agronomic characters. High heritability in the
broad-sense had also been found for total glycoalkaloid
content in tetraploid potatoes in a study by Sanford and
Sinden (1972).
Heat tolerance is not necessarily correlated with high
levels of glycoalkaloid in tubers and thus it is possible
to find tolerant clones with low content of glycoalkaloid
when grown under conditions of high temperatures
(Veilleux et al. 1997).
Potato breeding in the past was mainly focused on
tuber yield, quality and resistance to pests and diseases,
quality, being less oriented towards tolerance to abiotic
stress (Schafleitner et al. 2011). Breeding for heat
tolerance should focus on effect of high temperatures
on tuberization, since potato tuber initiation and
development are very sensitive to high temperatures
(Muthoni and Kabira 2015).
Since 2004, The International Potato Center (CIP)
has sought to improve the heat tolerance of its late-blight
resistant population, B3, by developing the new late blight
and temperature tolerant ‘LBHT’ population (Gastelo
et al. 2015). The aim was to obtain potato clones with
high levels of resistance to late blight, with high tuber
yield under high temperatures, more than 20°C at night,
Table 1: Experimental sites in Perú for narrow-sense and broad-sense heritability for yield components and glycoalkaloid content
Site Altitude masl Latitude Longitude Growing season
San Ramon 828 11°08’S 75°20’W July- October 2014
La Molina 244 12°05’S 76°56’W November 2014 – February 2015 Tropical Lowlands
Majes 1294 16°28’S 72°06’W November 2014 – February 2015 Tropical mid-elevations
low glycoalcaloid content and early maturity (90 days),
adapted to tropical mid-elevation environments.
In the present study, we determined the narrow-sense
heritability of yield components and broad-sense heritability
of glycoalkaloid content and its correlation with tuber
yield in ‘CIP’s LBHT’ population. The aim was to use yield
components as secondary traits for improving tuber yield and
heat tolerance in potato breeding, ensuring the productivity
and quality of new heat-tolerant potato varieties, with high
tuber yield and low glycoalkaloid content even when raised
under warm environments. These progenitors could then
serve as parents in breeding programs.
2 Methods
2.1 Estimation of narrow-sense heritability
for yield components
In order to determine the narrow-sense heritability for
yield components in the LBHT population, 32 clones were
taken at random and were crossed at the Huancayo, CIP–
Experimental station in Peru in 2012 under greenhouse
conditions using a North Carolina II mating design, in
four sets with four female and four male progenitors,
producing 16 full-sib progenies per set, in total 64 full-sib
families. Tuber families of each progeny were generated
in 2013 in Lima under greenhouse and then increased in
field conditions at Huancayo in November 2013 and April
2014. During 2014 - 2015, we evaluated the progenies
under field conditions, in three warm environments
in Peru: San Ramon, planted in July and harvested in
October 2014, La Molina, and Majes, planted in November
2014 and harvested in February 2015 (Table 1). Average
temperatures at these sites at night were between 16 to
23°C, and daytime temperatures fluctuated between 23 to
28°C. Temperature is very important for the formation of
potato tubers, especially nighttime temperatures, since
the night-time process of tuberization is inhibited above
20°C, (Figure 1,2,3,4).
Agroecological zone T° Average °C
Day Night
Humid tropical mid- 28 23
elevations
23 20
23 16
Unauthenticated
Download Date | 11/19/17 1:06 AM
  Heritability for Yield and Glycoalkaloid Content in Potato Breeding under Warm Environments   563

Figure 1: Temperatures during growing season in Majes, Peru, November 2014 to February 2015

Figure 2: Temperatures during growing season in San Ramon, Peru 2015

We used randomized complete block (RCB) design
with four sets and 3 replications of 50 genotypes per
progeny. The plots consisted of two rows of 25 plants
each, the distance between rows was 0.90 m and
between plants 0.30 m. The field management was the
same as in commercial potato fields with sowing and
harvesting done manually. No fungicides were applied.
Irrigation was achieved by row in La Molina and San
Ramon and by drip irrigation in Majes. Vine-killing
occurred at 90 days after planting and the harvest
took place one week later. Number of harvested plants,
number and weight of marketable and non-marketable
tubers were recorded. Additive and non-additive genetic
variance and narrow-sense heritability were calculated
from the male and female variance components,
assuming random effects of parents and environments,
Model II, (Tables 2,3) (Hallauer and Miranda 1981). The
correlation between yield components and tuber yield
was calculated using Pearson’s correlation coefficient
(p<0.01).

Unauthenticated
Download Date | 11/19/17 1:06 AM
564   M.A. Gastelo Benavides, et al.

Figure 3: Temperatures during growing season in La Molina, Peru, November 2014 to February 2015

Figure 4: Average day and night temperatures during the growing season in San Ramon 2014

2.2 Estimation of broad-sense heritability
for Glycoalkaloid content
We also estimated, the broad-sense heritability for
accumulation of glycoalkaloid content in tubers of LBHT
clones with resistance to Late Blight and heat tolerance
under high temperature conditions. In 2014-2015, three
experiments were conducted under warm environments:
San Ramon, La Molina and Majes in Peru (Table 1).
Twenty LBHT elite clones (Centro Internacional de la
Papa 2014), and control varieties: Unica and Desiree,
were evaluated in a RCB design with three replications

Unauthenticated
Download Date | 11/19/17 1:06 AM
  Heritability for Yield and Glycoalkaloid Content in Potato Breeding under Warm Environments   565

Table 2: Analysis of variance of Mating Design II North Caroline, over environments for model II
Source of variation df Mean Squares Expected mean squares

Environment (E) e-1

Sets (S) s-1
ExS (e-1)(s-1)
Replication/E/S se(r-1)
Male/S s(m-1) M7 σ²e + rσ²fme + rfσ²me + reσ²fm + rfeσ²m
Female/S s(f-1) M6 σ²e + rσ²fme + rmσ²fe + reσ²fm + remσ²f
Male x female/S s(m-1)(f-1) M5 σ²e + rσ²fme + reσ²fm
Male/S x E s(m-1)(e-1) M4 σ²e + rσ²fme + rfrσ²me
Female/S x E s(f-1)(e-1) M3 σ²e + rσ²fme + rmσ²fe
Male x Female/S x E s(m-1)(e-1)(f-1) M2 σ²e + rσ²fme
Pooled Error es(r-1)(mf-1) M1 σ²e
Total smfer-1

Table 3: Estimates of Variance Components in Mating Design II over environments for model II for male and female components
Variance Components Male Female

Error Variance σ²e = M1 σ²e = M1

Variance of Male x σ²mfe = (M2-M1)/r σ²mfe = (M2-M1)/r
female x Environment
Variance Male or σ²me = (M4-M2)/rf σ²fe = (M3-M2)/rm
Female x Environment
Variance Male x σ²mf = (M5-M2)/re σ²mf = (M5-M2)/re
Female
Variance Male or σ²m = (M7-M5-M4+M2)rfe σ²f = (M6-M5-M3+M2)rme
Female
Variance Additive σ²A = 4σ²m σ²A = 4σ²f
Variance Non-Additive σ²NA = 6σ²mfe/fe + 4σ²me/e + 6σ²mf/f σ²NA = 6σ²mfe/me + 4σ²me/e + 6σ²mf/m
Phenotypic Variance σ²phenotypic = σ²A + σ²NA + σ²e/rfe σ²phenotypic = = σ²A + σ²NA + σ²e/rfe
Heritability h² = 4σ²m /(σ²e/rfe + 6σ²mfe/fe + 4σ²me/e + 6σ mf/f + 4σ²m) h² = 4σ²f /(σ²e/rme + 6σ²mfe/me + 4σ²me/e + 6σ mf/m + 4σ²f )

of 20 plants each (Table 4). Spacing of rows in the plots
was 0.9m, each row containing 10 plants spaced at 0.3m.
Field management was commercially based with manual
operations throughout. Vines were killed 90 days after
planting and the harvest was made one week later.
Fungicides were not applied for the control of late blight.
In all experiments, the number of harvested plants, and
number and weight of marketable and non-marketable
tubers were recorded. For glycolakaloid analysis, in
all locations, 15 peeled tubers of medium and uniform
size, free of greenings per clone per each replicate were
sampled and then freeze-dried and milled from each
clone were prepared and stored at -80°C until analysis of
total glycoalkaloids was performed. Total glycoalkaloid
was analysed using the method described by Hellenas et
al. (1986). Briefly, glycoalkaloid extraction was achieved
using methanol and chloroform before concentration at
60°C in a rotary evaporator. The extract was transferred
to a 2% acetic acid solution and then purified using
ammonium hydroxide at 85 °C and ultracentrifugation
at 27,000 rpm. The pellet was reacted with 85% ortho-
phosphoric acid and read at 408 nm. The determination
of total glycoalkaloids was achieved against a standard
curve of α-chaconine as reference. Heritability in the
broad-sense at 95% of confidentiality (Knapp 1986) and
Pearson’s correlation (p<0.01) between tuber yield and
glycoalkaloid content were estimated.
In both experiments, the GLM procedure in SAS for
Windows, version 9.4 (SAS Institute Inc.) software was
used for statistical and genetic analysis.
3 Results
3.1 Estimation of narrow-sense heritability
for Yield components
A combined analysis of variance for tuber number,
yield per plant and average tuber weight showed highly
significant differences (p<0.01) for environment, male
and female/sets, and male/set x environment. Female/set
x environment was not significant for tuber number per

Unauthenticated
Download Date | 11/19/17 1:06 AM
566   M.A. Gastelo Benavides, et al.

Table 4: Clones from LBHT, evaluated for glycoalkaloid content in San Ramon, La Molina and Majes in 2014-2015

Clone Female Male Resistance Tolerance

LB PVX PVY Heat Drought

CIP302506.39 Bzura CIP393280.57 R ER ER T NT

CIP302533.38 CIP393371.159 CIP396272.43 R ER ER T NT
CIP302533.49 CIP393371.159 CIP396272.43 R ER ER T NT
CIP393371.58 CIP387170.16 CIP389746.2 R ER S T NT
CIP398098.203 CIP393371.58 CIP392639.31 R S ER T T
CIP398098.204 CIP393371.58 CIP392639.31 R S ER T NT
CIP398098.570 CIP393371.58 CIP392639.31 R ER S T NT
CIP398180.289 CIP392657.171 CIP392633.64 R ER S T T
CIP398180.292 CIP392657.171 CIP392633.64 R ER S T T
CIP398190.200 CIP393077.54 CIP392639.2 R S S T NT
CIP398190.605 CIP393077.54 CIP392639.2 R S S T NT
CIP398190.615 CIP393077.54 CIP392639.2 R ER S T NT
CIP398192.213 CIP393077.54 CIP392633.54 MR ER S T T
CIP398192.553 CIP393077.54 CIP392633.54 R ER S T NT
CIP398193.158 CIP393077.54 CIP392633.64 R ER S T NT
CIP398201.510 CIP393242.5 CIP392633.64 R ER ER T NT
CIP398208.219 CIP393371.58 CIP392633.64 R ER ER T T
CIP398208.505 CIP393371.58 CIP392633.64 R ER ER T NT
CIP398208.620 CIP393371.58 CIP392633.64 R ER S T NT
CIP398208.670 CIP393371.58 CIP392633.64 R ER ER T NT
R = Resistant; MR = Moderate resistance; ER = Extreme resistance; S = Susceptible; T = Tolerant; NT= No tolerant ; LB = Late blight;
PVX =Potato Virus X; PVY = Potato Virus Y

Table 5: Analysis of variance from North Carolina mating design II over environments for yield components, model II under warm environ-
ments 2014-2015
Source of variation df Tuber number per plant Tuber yield per plant (g) Average tuber weight (g)

Environment (E) 2 1535.62 ** 22.32 ** 80411.32 **

Sets (S) 3 11.71 ns 0.20 ** 4676.97 **
ExS 6 56.11 ** 0.45 ** 1732.40 **
Replication/E/S 24 3.94 ns 0.02 ns 210.34 ns
Male/S 12 M7 31.55 ** 0.14 ** 4463.27 **
Female/S 12 M6 27.99 ** 0.11 ** 839.12 **
Male x Female/S 36 M5 8.99 ** 0.07 ** 485.04 **
Male/S x E 24 M4 16.36 ** 0.07 ** 553.80 **
Female/S x E 24 M3 7.36 ns 0.100 ** 467.86 **
Male x Female/S x E 72 M2 9.63 ** 0.07 ** 333.67 **
Pooled Error 360 M1 3.64 0.01 159.78
CV(%)   20.77 17.63 18.83

CV, coefficient of variation    

**, significant statistical differences at P<0.01    

plant, but highly significant for average tuber weight and
tuber yield per plant (p<0.01) (Table 5).
For tuber number per plant, additive genetic variance
was higher than non-additive genetic variance estimated
from male and female variance component; for average
tuber weight additive genetic variance was higher than
non-additive genetic variance, estimated from male
variance component, conversely non-additive genetic
variance was higher than additive genetic variance
estimated from female variance component. For tuber
yield per plant additive and non-additive genetic
variances had similar magnitudes estimated from male
variance component and non-additive genetic variance
was higher than additive genetic variance from female
variance components, (Table 6).
Narrow-sense heritability of tuber number per plant
estimated from both male and female variance was high
h²=0.50 and 0.76 respectively, for average tuber weight
estimated from male variance component was very high
h²=0.83, but estimated from female variance component

Unauthenticated
Download Date | 11/19/17 1:06 AM
  Heritability for Yield and Glycoalkaloid Content in Potato Breeding under Warm Environments   567

Table 6: Estimates of additive and non-additive genetic variance, and Narrow-sense heritability, estimated from male and female variance
components under warm environments

Component σ²A σ²NA σ²Phenotypic h²

Estimated from male variance components

Tuber number per plant 1.759 1.640 3.500 0.50 ± 0.14
Average tuber weight (g) 417.567 78.668 500.673 0.83 ± 0.13
Tuber yield per plant (g) 0.007 0.010 0.017 0.41 ± 0.12

Estimated from female variance components

Tuber number per plant 2.363 0.640 3.104 0.76 ± 0.14
Average tuber weight (g) 24.434 69.118 97.991 0.25 ± 0.14
Tuber yield per plant (g) 0.005 0.012 0.017 0.30 ± 0.13
h2 = heritability; σ²A = additive genetic variance; σ2NA = non-additive genetic variance.

Table 7: Pearson correlations between tuber yield and its components in 64 full-sib families evaluated under warm environments. 2014-2015

  Tuber Yield/Plant R² Number of tubers/ R² Average tuber weight

plant

Tuber Yield/Plant (g plant-1) 1        

Number of tubers per plant 0.769** 0.59 1    
Average tuber weight (g) 0.742** 0.55 0.213* 0.05 1

*Correlation is significant at P<0.05 and ** P<0.01

Table 8: Estimates of variance components and broad-sense heritability for glycoalkaloid content

Variance components Glycoalkaloid content in peeled tubers

(mg/100g FW)

Genetic variance σ²c 66.87

Genetic x Environment Variance σ²ce 114.87
Error Variance σ²e 6.79
Phenotypic Variance σ²p 105.91
Broad-sense Heritability h² 0.46 < 0.63 < 0.80

was low h²=0.25. The estimate for tuber yield per plant
from male h²=041 was higher than that estimated from
female component h²=0.30.
Pearson correlations among tuber number per plant
and average tuber weight were highly significant (p<0.01)
and positive correlation to tuber yield per plant r=0.769
and 0.742 respectively (Table 7). Tuber number per plant
and average tuber weight predict the tuber yield per plant
in R²=0.59 and 0.55 respectively.
3.2 Estimation of broad-sense heritability
for Glycoalkaloid content
Combined analysis of variance for marketable tuber yield
(tha-1) and glycoalkaloid content (mg/100g FW) in peeled
tubers showed highly significant differences among
environments, clones and clones x environments (p<0.01).
Estimates of genetic variances and broad-sense heritability
for glycoalkaloid content in peeled tubers are shown in
table 8, where broad-sense heritability of glycoalkaloid
content was high, h²=0.63, and Pearson’s correlation
between glycoalkaloid content and marketable tuber yield
under high temperatures in all locations was weak r=0.33
and R²=0.11 (p<0.01). Correlations per locality was also
weak, with r= 0.34, 0.26 and 0.09 and R²=0.12, 0.07 and
0.01 for La Molina, Majes and San Ramon respectively.
Marketable tuber yield under strong heat stress and
with added virus pressure in the humid tropical mid-
elevation environment of San Ramon was in the range
from 9.95 to 48.42 tha-1. compared to the average over three
warm environments of from 8.08 to 32.49 tha-1.
Glycoalkaloid content in San Ramon, best placed in
order to select clones with heat tolerance, ranged from
1.27 to 90.99 mg/100g FW. Ten clones showed high tuber
yield but lower glycoalkaloid content, lower that the safe

Unauthenticated
Download Date | 11/19/17 1:06 AM
568   M.A. Gastelo Benavides, et al.

level of glycoalkaloid in tubers (20 mg/100g FW). Eight of programs (Table 9, Figure 5). The level of glycoalkaloid
these clones showed concentrations of 1.27 to 7.52 mg/100g content in La Molina and Majes was lower than in San
FW. These values are within the range recommended by Ramon. In la Molina the range was from 1.06 to 28.03
Jacob van Dam (2002), for plants to be used in breeding mg/100g FW and in Majes from 0.77 to 29.32 mg/100g FW.

Table 9: Marketable tuber yield and glycoalkaloid content in 20 LBHT clones under warm environments. 2014- 2015
Glycoalkaloid mg/100g FW Marketable tuber yield tha-1

Clone San Ramon La Molina Majes San Ramon La Molina Majes Average

18.56 0.92 2.54 37.36 22.98 32.60 30.98

CIP302506.39
CIP302533.38 19.87 4.36 1.85 41.76 18.77 17.72 26.08
CIP302533.49 54.87 25.53 29.32 47.76 18.54 25.51 30.60
CIP393371.58 29.33 14.25 4.19 29.03 28.99 33.26 30.43
CIP398098.203 35.22 25.61 3.37 25.81 10.91 19.69 18.81
CIP398098.204 40.55 2.10 4.77 25.92 5.53 19.82 17.09
CIP398098.570 5.65 1.24 1.93 29.41 16.69 21.33 22.48
CIP398180.289 4.57 5.44 1.84 34.05 20.72 23.31 26.03
CIP398180.292 18.16 28.03 1.63 32.25 30.96 34.26 32.49
CIP398190.200 2.51 3.28 1.82 26.86 18.71 23.85 23.14
CIP398190.605 4.16 0.84 0.82 23.48 15.43 17.16 18.69
CIP398190.615 2.94 5.67 1.41 24.54 15.23 16.38 18.72
CIP398192.213 1.27 1.10 1.10 34.79 12.70 27.77 25.09
CIP398192.553 7.52 2.76 0.77 20.60 18.81 12.06 17.16
CIP398193.158 1.51 1.06 1.69 9.95 3.88 10.41 8.08
CIP398201.510 26.25 3.39 2.99 48.42 8.64 24.46 27.17
CIP398208.219 3.56 16.17 2.02 31.76 16.43 22.53 23.57
CIP398208.505 90.99 13.22 12.70 27.39 28.32 37.01 30.91
CIP398208.620 14.28 2.05 2.23 34.36 6.33 18.01 22.90
CIP398208.670 3.20 1.56 1.59 23.78 17.43 18.21 19.81
DLS (P<0.05) 20.15 3.97 2.98 9.16 4.30 3.09 3.46
C.V. % 21.70 29.95 30.59 17.60 18.58 9.06

Figure 5: Marketable tuber yield in ten clones with low glycoalkaloid content under warm environment. San Ramon 2014

Unauthenticated
Download Date | 11/19/17 1:06 AM
  Heritability for Yield and Glycoalkaloid Content in Potato Breeding under Warm Environments   569

4 Discussion allow us to select clones with high tuber yield and low
glycoalkaloid content.
Under humid tropical mid-elevation environments
4.1 Estimation of narrow-sense heritability
(San Ramon), ten LBHT clones showed high marketable
for yield components tuber yield and a glycoalkaloid content that was lower
than the safe level in tubers (20 mg/100g FW) as laid
Late-blight heat tolerant (LBHT) clones selected under
down by Jacob van Dam (2002). Eight clones, showed low
the humid tropical mid-elevation environment in San
glycoalkaloid content and high marketable tuber weight
Ramon, used in this study, showed adaptation to high
and so would be suitable parents in breeding programs
temperatures in warm environments, one of the major
aimed at improving heat tolerance with minimum risk
abiotic factors affecting the production of tubers in potato
of glycoalkaloid accumulation under high temperatures
crop under climate change.
stress or to be used to develop new varieties suitable for
Narrow-sense heritability for tuber yield per plant
hight heat stress (Table 9, Figure 2).
indicated that the estimated genetic and non-additive
In Tropical lowlands (La Molina) and tropical mid-
genetic variances from male and female variance
elevation (Majes) environments, the glycoalkaloid content
components are sufficient in order to improve the
was lower than that in the humid tropical mid-elevation
frequency of genes and genotypes favorable for this trait
(San Ramon), with its the lower temperatures, especially at
under warm environments and climate change conditions.
night during the tuberization season, demonstrating that
The narrow-sense heritability of the number of tubers
as night temperatures increase above 20°C glycoalkaloid
per plant was high for both male and female variance
levels can increase, as observed in San Ramon, where
components, the additive genetic variance being more
temperatures were higher (see Table 1,9 and Figures 1- 4).
important than the non-additive genetic variance, and
High heritability for glycoalkaloid concentration
allowing us to be sure of selecting clones producing more
and weak correlation with marketable tuber yield, will
tubers. The high association found with tuber yield per
allow us to select clones with high tuber yield and low
plant, where 59% depends on the number of tubers, will
glycoalkaloid content suitable as candidate varieties and/
allow us to select clones with high tuber yield using this
or parents in breeding programs.
trait as indirect selection criterion.
Given the our finding of high concentrations of
The narrow-sense heritability of average tuber weight
glycoalkaloid in population under warm environments,
was very high, when estimated from male variance, but
assessment of parental value for this feature is
was low when it was estimated from the female variance
recommended when selecting clones for further breeding.
component, possibly due to maternal cytoplasmic effects
affecting the expression of this character and a property
Acknowledgments: This research was undertaken as
that should be further researched. In this study, average
part of the CGIAR Research Program on Roots, Tubers and
tuber weight was highly correlated with and thus accounts
Bananas (RTB). It has also received financial support from
for 55% of tuber yield. This agrees with Jefferies and
USAID.
MacKerron (1987), who found that average tuber weight
was an important factor influencing tuber yield, and
Thompson et al. (1983) who had suggested using average References
tuber weight as an indirect selection criterion.
High heritability for yield components in the LBHT Birch P.R.J., Bryan G., Fenton B., Gilroy E., Hein I., Jones J.T., Prashar
population indicates that there is enough genetic A., Taylor M.A., Torrance L., Toth I.K., Crops that feed the
world: Potato: are the trends of increased global production
variability for further gains in selectively breeding for
sustainable? Food Security, 2012, 4, 477-508
tuber yield under warm conditions. Cabello R., Monneveux P., Bonierbale M., Khan A., Heritability of
yield components under irrigated and drought conditions in
andigenum potatoes. Am. J. of Potato Res., 2014, 91, 492-499
4.2 Estimation of broad-sense heritability Centro Internacional de la Papa (International Potato Center),
Catalogue of potato varieties and advanced clones, 2014,
for glycoalkaloid content http://www.cipotato.org/catalogue
Ewing E.E., Heat stress and the tuberization stimulus. Am. Potato J.,
The second study estimating the broad-sense heritability 1981, 58 31-49
of glycoalkaloid content, indicated high heritability and
a weak correlation with marketable tuber yield. This will

Unauthenticated
Download Date | 11/19/17 1:06 AM
570   M.A. Gastelo Benavides, et al.
Friedman M., Potato glycoalkaloids and metabolites: Roles in
the plant and in the Diet. J. Agric. Food Chem., 2006, 54,
8655-8681
Gandolfi Flávio R.; Pereira C.A., Genetic gains for heat tolerance in
potato in three cycles of recurrent selection. Crop Breeding
Appl. Biotechnology, 2011, 11(2), http://dx.doi.org/10.1590/
S1984-70332011000200005   M., Adaptation of the potato crop to changing climates. In crop
Gastelo M., Diaz L., Landeo J.A., Bonierbale M., New elite potato
clones with heat tolerance, late blight and virus resistance to
address climate change. In: Low, J.; Nyongesa M., Quinn S.,
Parker M. (eds). Potato and sweetpotato in Africa. Transforming
the value chains for food and nutrition security. Oxfordshire
(UK). CABI International. ISBN 978-1-78064-420-2, 2015, pp.
143-152, http://dx.doi.org/10.1079/9781780644202.0143
http://hdl.handle.net/10568/72563
Hallauer A.R., Miranda J.B., Quantitative genetics in maize breeding.
The Iowa State University Press, Ames, Iowa, USA. 1981
Hellenas K.E., A simplified procedure for quantification of potato
glycoalkaloids in tubers extracts by HPLC: Comparison with
ELISA and colorimetric method. J. Sci. Food Agric., 1986, 37,
776-782
Hijmans R., The effect of climate change on global potato
production. Am. J. of Potato Res., 2003, 80, 271-280
Jefferies R.A., MacKerron D.K.L., Aspects of the physiological basis
of cultivar differences in yield of potato under drought and
irrigated conditions. Potato research, 1987, 30, 201-217
Knapp S.J., Confidence intervals for heritability for two-factor design
single environment linear models. Thor. Appl. Genet., 1986, 72,
587-591
Levy D., Veilleux R., Adaptation of potato to high temperatures and
salinity. A review. Am. J. of Potato Res., 2007, 84, 487-506
Luthra SK, Kamlesh Malik, Gupta V.K., Singh B.P., Evaluation of
potato genotypes under high temperature stress conditions.
Crop Improv. 2013, 40(1), 74-80
Mondy N.I., Ponnapalam Y., Determination of total glycoalkaloids
(TGA) in dehydrated potatoes. J. Food Sci., 1983, 48(2), 612-614
Muthoni J., Kabira J.N., Potato production in the hot tropical areas
of Africa: Progress made in breeding for heat tolerance. J. of
Ag.Sci., 2015, 7(9), 220-227
Rykaczewska K., The effect of high temperatures occurring in
subsequent stages of plant development on potato yield and
tuber physiological defects. Am. J. of Potato Res., 2015, 92,
39-349
Sanford L.L., Sinden S.L, Inheritance of potato glycoalkaloid. Am.
Potato J., 1972, 49, 209- 217
SAS Institute. Copyright©, Base SAS® 9.4 Procedures Guide:
Statistical Procedures, Second Edition. SAS Institute Inc., Cary,
NC, USA, 2013
Schafleitner R., Ramirez J., Jarvis A., Evers D., Gutierrez R., Scurrah
adaptation to climate change. First Edition, Chapter 11. John
Wiley & Sons. Ltd. Published by Blackwell Publishing Ltd., 2011
Smith D.B., Roddick J.G., Jones J.L., Potato glycoalkaloids: Some
unanswered questions. Trends in Food Science & Technology,
1996, 7, 126-131
Storey R.M.J., Davies H.V., Tuber quality In: P. Harris ED. The potato
crop. 2nd ed. Chapman & Hall, London, 1992, p. 507-569
Thompson P.G., Mendoza H.A., Plaisted R.L., Estimation of genetic
parameters for characters selected for potato propagation by
true seed in an andigena population. American Potato Journal,
1983, 60, 393-401
Van Dam J., Genetic characterization of agronomic and morpho-
logical traits and the development of DNA markers associated
with total glicoalkaloid content in the tubers of the tetraploid
potato (Solanum tuberosum), PhD Thesis Wageningen
University, 2002, pp.101
Van Dam J., Levin I., Struik P.C., Levy D., Genetic Characterisation of
tetraploid potato (Solanum tuberosum L.) emphasizing genetic
control of total glycoalkaloid content in the tubers. Euphytica,
1999, 110, 67-76
Veilleux R.E., Paz M.M., Levy D., Potato germoplasm development
for warm climates: genotypic enhancement of tolerance to heat
stress. Euphytica, 1997, 98, 83-92
Zheng S-L., Wang L-J., Wan N-X., Zhong L., Zhou S-M., He W., Yuan
J-C., Response of potato tuber number and spatial distribution
to plant density in different growing seasons in Southwest
China, Frontiers in Plant Science, 2016, 7, 365
Yuen J., Forges G., Estimating the level of susceptibility to
Phytophthora infestans in potato genotypes. The American
Phytopathological Society, Phytopathology, 2009, 99(6),
782-786
Unauthenticated
Download Date | 11/19/17 1:06 AM