Euphytica (2018) 214:116

https://doi.org/10.1007/s10681-018-2201-8

Combining ability analysis of earliness and yield of potato

(Solanum tuberosum L.) genotypes in Uganda
Prossy Namugga . Julia Sibiya . Rob Melis . Alex Barekye

Received: 5 January 2018 / Accepted: 18 June 2018 / Published online: 21 June 2018
Ó Springer Nature B.V. 2018
Falta a justificativa do trabalho
Abstract Potato (Solanum tuberosum L.) is a major and 0.78 for days to 50% flowering. The predomi-
food and cash crop mainly grown by small-scale nance of additive genetic effects imply that, genetic
farmers in the highland regions of Uganda. Changing gains can be achieved through different selection
global weather patterns require varieties that are able methods and traits transferred to the respective
to grow within the short rainfall cycles and yield progenies. Parents Rwangume, 396,038.107,
optimally under the prevailing conditions. The objec- 395,011.2, NKRK19.17, 393,077.54, Kimuri, and
tives of this study were to estimate the combining 392,657.8 had desirable GCA effects for the number
ability effects for early maturity, yield and yield of days to flowering and yield related traits. Families
related traits in potato. Eighteen F1 families generated of Rwangume 9 NKRK19.17, 393,077.54 9 395,011.2,
from two sets of 12 parents using a North Carolina 396,038.107 9 Rwangume and 396,038.107 9
Design II were evaluated for days to 50% flowering, 395,011.2 had desirable SCA effects for yield and
leaf senescence, yield and yield related traits in two number of days to 50% flowering. The selected parents
different locations. Both additive and non-additive and families will be subjected to further clonal
genetic effects influenced the expression of traits. evaluation and selection.
However, additive genetic effects were predominant
over the non-additive for most of the traits. The GCA/ Keywords Combining ability  Earliness  Gene
SCA ratios were 0.68 and 0.78 for days to 50% action  Heritability  Maturity  Solanum tuberosum
flowering and average tuber weight. Broad sense
heritability estimates were 0.70 for total tuber weight Poderia falar melhor da importância desse alimente
para países como a Uganda.
Introduction
P. Namugga (&)  J. Sibiya  R. Melis
African Centre for Crop Improvement, School of Potato (Solanum tuberosum L.) is a major food and
Agricultural, Earth and Environmental Sciences, cash crop, mainly grown by small-scale farmers in the
University of KwaZulu-Natal,
highland regions of many African countries. Uganda is
Private Bag X01, Scottsville, Pietermaritzburg 3209,
South Africa the ninth largest producer of potato in Africa with an
e-mail: [email protected] annual production of 188,000 tons harvested from
about 39,000 ha per year (FAOSTAT 2016). Potato
P. Namugga  A. Barekye
yields in Uganda have remained low about 4.8 t ha-1
National Agricultural Research Organisation (NARO),
Kachwekano Zonal Agricultural Research and (FAOSTAT 2016) against a potential of about
Development Institute, P.O. Box 421, Kabale, Uganda 25 t ha-1 which can be achieved under good

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management and when suitable varieties are deployed. limited. This is accompanied by the growing need for
These low yields have been attributed to a number of early maturing and high yielding varieties to counter-
confounding factors that are biotic, abiotic, and socio- act the effects of changing weather patterns. Earliness
economic constraints, as well as poorly adapted and is a quantitative trait that is affected by genetic and
adopted varieties. The most preferred attributes by physiological factors of a plant besides environmental
most potato farmers in Uganda were identified as high conditions (Basbag et al. 2007). Earliness is reported
yields followed by marketability, resistance to blight to contribute to disease escape especially to infection
and early maturity (Namugga et al. 2017a, b). pressures that appear late in the season. Earliness is
Crop maturity is an agronomic trait where crops also beneficial in areas with multiple cropping
undergo progressive growth stages from emergence to systems, limited land or short favourable growing
senescence, characterized by their reproductive capac- seasons. In Uganda, potato is a smallholder farmer
ity and phenology (Khan et al. 2013). Potato maturity crop so early maturing varieties would allow favour-
is normally assessed by monitoring vine characteris- able rotation patterns and contribute greatly to food
tics and change in colour of the potato plant leaves is security. This study was undertaken to determine the
an indicator of crop maturity (Haga et al. 2012; combining ability effects for early maturity, yield and
Iragaba 2013). Potato varieties are classified into yield related traits in potato in order to develop potato
maturity types based on the lengths of the season clones that are early maturing with high yield
required to produce a harvestable product (Haga et al. potential.
2012; Khan et al. 2013). Variability among varieties
for the number of days from planting to maturity has
led to designation of the potato varieties as early, Materials and methods
medium, late and very late maturity classes (Ruzukas
et al. 2009). Breeders normally evaluate maturity Parental materials and crosses
Método Carolina do norte II
period while developing new varieties because it is a
critical aspect in commercial potato production. Twelve genetically diverse clones were selected and
Standard maturity measurements in potato are lacking used as parents. Six clones were obtained from the
because tubers are produced underground and moni- National Potato Program of Uganda and four were
toring their development presents many challenges. advanced clones from the International Potato Centre
As a result, breeders commonly assess potato maturity (CIP) belonging to population B3C2 with variable
classes based on physiological changes in the potato resistance to late blight. These materials were selected
vine. Tuber production is associated with changes at based on their flowering abilities, number of days to
the whole plant level, such as reduction in leaf flowering, flowering duration and medium to high
development, flowering, fruit set and change of foliar yields (Namugga et al. 2017a, b). Parents were
colour (Haga et al. 2012). assembled into two sets of six parents each based on
In potato, both the general combining ability their flowering abilities, yield and resistance to late
(GCA) effects of parents and specific combining blight. Crosses were made using a North Carolina
ability (SCA) effects of their progeny are important in Design II (NCD II) to generate 18 families (Table 1).
determining economic traits. This is due to the fact all In the first set three female clones (Rwangume,
genetic effects are fixed at the F1 generation through 396,026.103 and 396,038.107) were crossed with
the development of clones without further segregation three males (Kimuri, 391,046.14 and NKRK19.17).
(Muthoni et al. 2012). Earliness in potato has not In the second set, three female parents; 392,657.8,
received sufficient attention in Uganda and elsewhere 393,220.54 and 396,038.107 were crossed with three
as there seems to be limited information regarding the males; NKRN59.48, Rwangume and 395,011.2. Par-
subject. However, some findings suggest that both ents Rwangume and 396,038.107 were used in both
additive and non-additive genetic effects control sets. Controlled hand pollination was performed at
earliness in potato, though additive genetic effects flowering following emasculation (Acquaah 2007). At
were found to be more important (Iragaba 2013). maturity, berries of the same cross were harvested and
However, detailed information regarding inheritance bulked. In total, 18 families were generated (two sets
of this trait and combining ability of parents is still of nine families each).

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Table 1 Description of Set Parent Parent type Yield (t ha-1) Days to flowering
potato parents used in the
crossing block. Source: 1 Rwangume Female 25.2 59
Namugga et al. (2017a, b)
1 396,026.103 Female 33.1 54
1 396,038.107 Female 42.8 61
1 Kimuri Male 17.1 52
1 391,046.14 Male 34.6 52
1 NKRK19.17 Male 37.0 61
2 392,657.8 Female 43.7 55
2 393,077.54 Female 43.2 59
2 396,038.107 Female 42.8 61
2 Rwangume Male 25.2 59
2 395,011.2 Male 25.5 54
2 NKRN59.48 Male 30.0 61

Planting sites as the number of days from flowering to 100% of the

leaves dying off. At harvest, yield related data was
The field trials were established at Kachwekano taken in a plot for, total weight of all the tubers
research station and on a farm in Nyabumba. Both harvested in a plot (TTW) and expressed in t ha-1.
sites are located in South-western Uganda; Kach- Average tuber weight (ATW) was calculated as the
wekano is located in, 01°160 S 29°570 E at 2200 meters total tuber weight per plant divided by the total tuber
above sea level while Nyabumba at 2230 meters above number of tubers per plant.
sea level. Both sites have a bi-modal rainfall pattern
separated by a dry spell ranging from 30 to 60 days. Data analyses
Planting was done in March and harvesting in July
2017. Analysis of variance

Experimental design and trial establishment Data for the different traits over two sets and across
environments were subjected to the standard analysis
Experiments were established during the planting of variance using the using general linear model
season between March and June 2017 using a 7 9 4 (GLM) procedure of SAS 9.3 (SAS 2011) statistical
alpha lattice design with two replications at both program. Analyses of variance of NCD II pooled over
locations. Parents Rwangume and 396,038.107 were sets and across environments (Hallauer and Miranda
used in both set. The 18 F1 families were planted in 1988) were conducted for all the traits. Main effects
two row plots of 20 tubers each at 75 cm between rows due to female and male effects within sets are
and 30 cm between plants. Planting was done by hand independent estimates of GCA while male 9 female
and N:P:K: 17:17:17% fertilizer was applied at interaction effects represent SCA variance within sets.
100 kg ha-1. Pest and disease control was done using Spearman correlation coefficients were calculated for
recommended insecticides and fungicides. Crop hand the studied traits to determine their association.
weeding and ridging were carried out as
recommended. Estimation of general and specific combining ability
effects
Data collection
Data were analysed over sets and across environment.
Data were collected on days to flowering and leaf Parents were considered as fixed effects in the test of
senescence. Days to flowering was recorded as significance. The GCA and SCA values for each trait
number of days from planting to 50% flowering of were calculated following the NCII mating design
all the plants in a plot. Leaf senescence was measured across sites (Hallauer et al. 1988). General and specific

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combining abilities (GCA and SCA respectively) for the expression of all the traits (Table 2). The additive/
the parents and progenies were determined; and their non-additive genetic effects using Baker’s ratio was
significance tested (Singh and Chaudhary 1979). The highest for ATW (0.78) and lowest for LS (0.41).
relative importance of GCA and SCA in influencing
the performance of the crosses were estimated using Heritability estimates
the general predicted ratio (GPR) for all the traits
(Baker 1978). The broad sense heritability values were highest for
DAF (0.78) and lowest for ATW (0.39) (Table 3). The
Estimating heritability narrow sense heritability estimates were highest for
DAF (0.40) and the lowest for LS (0.09).
Heritability estimates were calculated using the
female additive variance for both narrow and broad General combining ability effects of parents
sense heritability (Dabholkar 1999) as follows: narrow
sense heritability; h2f = 4r2f /(r2e /r ? 4r2mf ? 4r2f )- Significant GCA effects were observed among all the
= VAf/VP; broad sense heritability; h2f = 4r2- f parents for ATW (Table 4). Female parents 392,657.8
? 4r2mf/(r2e /r ? 4r2mf ? 4r2f ) = VGf/VP. Where and 396,038.107 had significant GCA effects for DAF,
r = number of replication; r2e = environmental vari- while male parents Rwangume and NKRN59.48 had
ance = MSe; r2f = variance of female parents = GCAf GCA effects significant for TTW (Table 4). Among
variance = MSf; r2fm = variance due to interaction the female parents, 396,038.107 had the lowest GCA
between females and males = SCA variance = MSmf; effects for DAF (- 1.44) followed by Rwangume
VAf = additive genetic variance due to female parents; (- 0.42), while 392,657.8 had the highest (1.14). Male
VP = phenotypic variance; VGf = total genetic parents 395,011.2 and NKK 19.17 had the lowest
variance. GCA effects for DAF (- 0.59 and - 0.38), respec-
tively (Table 4). For yield and related traits, Rwan-
gume had the highest GCA effects for TTW (0.41)
Results followed by NKRK19.17 (0.28), while the lowest
GCA effect was for NKRN59.48 (- 0.20). Addition-
Analysis of variance for crosses across sites ally, the GCA effects for ATW were highest for
393,077.54 (0.02).
The combined analysis of variance and ratio of GCA/
SCA for days to flowering (DAF), leaf senescence Specific combining ability effects of families
(LS), total tuber weight (TTW) per hectare and
average tuber weight (ATW) per hectare among the Crosses largely showed significant SCA effects for ATW
families is presented in Table 2. Significant differ- while no significant SCA effects were observed for other
ences within sets observed for ATW (P B 0.05). The traits (Table 5). The highest SCA effects for TTW were
environmental effect was highly significant (P among the crosses of Rwangume 9 NKRK19.17 (0.36)
B 0.001) for all the traits except for number of days followed by 393,077.54 9 395,011.2 (0.25). Crosses of
to 50% flowering. The GCA mean squares for females 396,038.107 9 Rwangume had highest SCA effects for
(GCAf) were significantly different for DAF, and ATW (0.021). For number of days to 50% flowering,
TTW at P B 0.001, and P B 0.05, respectively. The families of Rwangume 9 NKRK19.17 (- 1.38) and
GCA mean square for males (GCAm) were signifi- 396,038.107 9 395,011.2 (- 0.75) had the lowest SCA
cantly different for DAF (P B 0.001) and TTW effects (Table 5).
(P B 0.05). The SCA effects for the crosses were
significant for all the tested traits except ATW. The Family means across locations
environmental interactions with families were only
significant for TTW. The GCAf effects were higher The mean number of days to flowering was 54, while
than male GCA effects for LS, TTW and ATW, while total tuber weight was 9.3 t ha-1. Average tuber
the additive male effects were slightly more for DAF. weight was 0.5 (Table 6). The number of days to
Overall, the GCA was more important than the SCA in flowering ranged from 44.1 to 62 days. Families with

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Table 2 Combined analysis of variance for days to 50% flowering, leaf senescence, and tuber yield and related traits evaluated at
two locations in Uganda in 2017
Source of variation df Mean square
DAF LS TTW ATW

Seta 1 29.73 13.91 6.63 0.12*

Site 1 39.15 1702.85*** 1387.66*** 1.64***
Replication (site) 2 71.49* 76.15 28.96* 0.00
Female (set) 10 81.68*** 40.4 22.26* 0.05
Male (set) 10 87.31*** 21.93 20.60* 0.02
Female 9 male (set) 20 79.45*** 89.75* 31.46** 0.02
Site 9 set 1 57.18 25.54 7.27 0.08
Site 9 female (set) 10 11.63 18.82 8.66 0.04
Site 9 male (set) 10 20.16 59.23 11.26 0.03
Site 9 female 9 male (set) 20 18.69 52.85 19.42* 0.02
Error 20.09 29.03 9.65 0.03
R-square 0.84 0.78 0.92 0.82
GCA/SCA ratio 0.68 0.41 0.58 0.78
a
Set set within an environment, df degrees of freedom, DAF days to 50% flowering, LS days to leaf senescence, TTW total tuber
weight, ATW average tuber weight, GCA/SCA ratio calculated according to Baker (1978)
*Significant at P B 0.05; **significant at P B 0.01; ***significant at P B 0.001

Table 3 Broad and narrow-sense heritability (H2 and h2) for positive and significant between LS and TTW (P
days to 50% flowering, leaf senescence, and tuber yield and B 0.001); LS and ATW (P B 0.01).
related traits evaluated at two locations in Uganda in 2017
Heritability/traits DAF LS TTW ATW
Discussion
H2 0.78 0.45 0.70 0.39
h2 0.40 0.09 0.28 0.34
The significant mean squares of families for days to
2 2
H broad sense heritability, h narrow sense heritability, DAF flowering and total tuber weight indicated the presence
days to 50% flowering, LS days to leaf senescence, TTW total of genetic variation among parents and their crosses.
tuber weight, ATW average tuber weight
This implies that genotypes that are early maturing
with high yields can be selected for. The significant
GCA and SCA mean squares of the traits observed
less days to flowering were 396,038.107 9 395,011.2 shows that both additive and non-additive gene action
(44.1) and 396,038.107 9 NKRN59.48 (47.5). The were involved in the expression of the traits.
average total tuber yield was 9.3 t ha-1 and families The Baker’s ratio for number of days to 50%
393,077.54 9 Rwangume (14.7 t ha-1) and (Rwan- flowering and leaf senescence were 0.68 and 0.41
gume 9 NKRK19.17 (13.0 t ha-1) were the best respectively. These results signify the predominance
yielders Average tuber weight ranged from 0.3 to 0.6 of additive genetic effects in controlling the number of
(Table 6). days to flowering and non-additive effects for leaf
senescence. These findings differ from what has been
Correlation between traits reported in other studies. For instance, Iragaba (2013)
found both additive and non-additive gene action
The correlations between the four traits across loca- control earliness in potato, though additive genetic
tions are presented in Table 7. Correlations were effects were more important. Buso et al. (2000)
reported both additive and non-additive effects to

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Table 4 Estimates of Traits DAF LS TTW ATW

general combining ability
(GCA) effects for days to Set one
50% flowering, leaf
Females
senescence, total tuber
weight and average tuber Rwangume - 0.421 - 0.403 - 0.238 - 0.002**
weight of 12 potato parents 396,026.103 0.319 - 0.045 0.188 0.002**
evaluated at two locations 396,038.107 0.102 0.448 0.049 0.000
in 2017
Males
Kimuri 0.298 - 0.602 - 0.031 0.003**
391,046.14 0.078 0.239 - 0.253 - 0.012**
NKRK19.17 - 0.376 0.362 0.284 0.009**
SE 0.900 1.680 0.270 0.001
Set two
Females
SE standard error, DAF 392,657.8 1.144** - 0.124 0.025 - 0.004**
days to 50% flowering, LS 393,077.54 0.299 - 0.382 - 0.008 0.018**
days to leaf senescence, 396,038.107 - 1.443** 0.506 - 0.016 - 0.014**
TTW total tuber weight,
Males
ATW average tuber weight
Rwangume 0.561 0.209 0.411* 0.002
*, **Significantly different
from zero at C 1.96 SE and 395,011.2 - 0.589 0.277 - 0.207 - 0.010
2.56 SE respectively NKRN59.48 0.028 - 0.486 - 0.200** 0.008
*significant at P B 0.05; SE 0.800 2.710 0.460 0.002
**significant at P B 0.01

Table 5 Estimates of Traits DAF LS TTW ATW

specific combining ability
(SCA) effects for days to Set one
50% flowering, leaf
Rwangume 9 Kimuri 0.296 0.215 - 0.040 - 0.003
senescence, total tuber
weight and average tuber Rwangume 9 391,046.14 1.079 0.249 - 0.318 0.007**
weight of 18 F1 potato Rwangume 9 NKRK19.17 - 1.375 - 0.465 0.359 - 0.004*
families evaluated at two 396,026.103 9 Kimuri - 0.507 0.295 0.190 0.002
locations in 2017
396,026.103 9 391,046.14 - 0.635 - 1.211 0.298 - 0.007**
396,026.103 9 NKRK19.17 1.142 0.916 - 0.488 0.004*
396,038.107 9 Kimuri 0.210 - 0.510 - 0.150 0.001
396,038.107 9 391,046.14 - 0.444 0.961 0.020 0.000
396,038.107 9 NKRK19.17 0.234 - 0.451 0.130 0.000
SE 2.670 5.030 0.800 0.002
Set two
392,657.8 9 Rwangume - 0.790 1.060 0.063 - 0.002
392,657.8 9 395,011.2 1.240 - 1.260 - 0.136 - 0.006**
SE standard error, DAF 392,657.8 9 NKRN59.48 - 0.450 0.210 0.073 0.008**
days to 50% flowering, LS 393,077.54 9 Rwangume - 0.480 0.420 0.185 - 0.019
days to leaf senescence, 393,077.54 9 395,011.2 - 0.490 0.450 0.254 0.017**
TTW total tuber weight,
393,077.54 9 NKRN59.48 0.970 - 0.870 - 0.439 0.001
ATW average tuber weight
396,038.107 9 Rwangume 1.260 - 1.470 - 0.249 0.021**
*, **Significantly different
from zero at C 1.96 SE and 396,038.107 9 395,011.2 - 0.750 0.810 - 0.118 - 0.011**
2.56 SE respectively 396,038.107 9 NKRN59.48 - 0.520 0.660 0.366 - 0.009**
*significant at P B 0.05; SE 2.410 8.140 0.060 0.007
**significant at P B 0.01

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Table 6 Family and parent Crosses DAF (days) LS (%) TTW (t ha-1) ATW (kg)
means of days to 50%
flowering, leaf senescence, Trait and mean performance across sites
tuber weight and average
Rwangume 9 Kimuri 56.3 28.5 11.8 0.4
tuber weight of 18 potato
families evaluated at two Rwangume 9 391,046.14 58.5 32.0 9.8 0.4
locations in 2017 Rwangume 9 NKRK19.17 46.9 29.6 14.6 0.5
396,026.103 9 Kimuri 56.0 30.3 14.4 0.5
396,026.103 9 391,046.14 54.6 32.4 14.0 0.4
396,026.103 9 NKRK19.17 59.9 36.6 13.0 0.5
396,038.107 9 Kimuri 58.0 29.0 12.5 0.5
396,038.107 9 391,046.14 54.5 38.3 12.3 0.4
396,038.107 9 NKRK19.17 55.4 33.9 14.9 0.5
392,657.8 9 Rwangume 58.9 35.1 14.4 0.5
392,657.8 9 395,011.2 62.4 26.1 11.1 0.4
392,657.8 9 NKRN59.48 58.1 28.9 12.0 0.5
393,077.54 9 Rwangume 56.8 31.5 14.7 0.5
393,077.54 9 395,011.2 52.1 31.9 12.5 0.6
393,077.54 9 NKRN59.48 60.4 23.6 9.8 0.6
396,038.107 9 Rwangume 56.8 27.5 13.0 0.5
396,038.107 9 395,011.2 44.1 36.9 11.0 0.3
396,038.107 9 NKRN59.48 47.5 33.3 8.7 0.4
Parents
Rwangume 55.0 34.8 16.1 0.6
396,026.103 45.3 24.5 7.9 0.3
396,038.107 56.4 31.6 7.1 0.5
Kimuri 53.5 37.3 7.3 0.3
391,046.14 57.5 32.0 6.1 0.5
NKRK19.17 45.8 25.3 6.1 0.4
392,657.8 46.5 20.3 9.4 0.5
393,077.54 54.0 30.0 12.8 0.6
395,011.2 56.8 20.5 14.4 0.8
DAF days to 50%
flowering, LS days to leaf NKRN59.48 58.3 35.3 8.5 0.5
senescence, TTW total tuber Mean 54.0 30.6 9.3 0.5
weight, ATW average tuber CV (%) 8.0 23.0 33.4 32.3
weight

influence maturity and total tuber yields. However, desirable GCA effects for days to flowering and passed
because traits are fixed in the F1 generation when these on to their progenies. Crosses of female parent
clones are developed, both genetic effects are of great 393,077.54 displayed significant and negative SCA
importance in potato breeding (Muthoni et al. 2012). effects for days to flowering and leaf senescence. The
According to Singh and Chaudhary (2007), parents best parents were 396,037.108, 393,077.54, Rwan-
with significant GCA and SCA effects in the right gume and NKRK19.17.
direction would be desired for the trait of interest. For The current study found that the GCA effects for
early maturity, the desirable gene action is negative, tuber yield related traits were more important than
while for yield, is positive. In this study, both desirable SCA effects. For example, the Baker’s ratio for total
and non-desirable GCA effects were passed on to the tuber weight was 0.58 and average tuber weight was
respective progenies. Parents Rwangume (- 0.42), (0.78). This implies that additive genes largely influ-
396,038.107 (- 1.44) and NKRK 19.17 (- 0.38) had enced the expression of these traits. The predominance

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tamanho
116 Page 8 of 9 Euphytica (2018) 214:116

Table 7 Phenotypic correlation between four traits of 18 studies (Muhinyuza et al. 2016; Hirut 2015). Simi-
potato families larly, Mehdi et al. (2008) found total tuber yield to be
Trait LS TTW DAF ATW largely influenced by higher number of tubers per
plant and tuber size. This denotes that improving one
Leaf senescence – trait would subsequently improve the other. The
Total tuber weight 0.343*** – significant positive association between leaf senes-
Days to flowering 0.139 0.131 – cence and days to 50% flowering, total tuber weight
Average tuber weight 0.278** 0.639*** 0.122 and average tuber weight obtained in this study, may
*Significant at P B 0.05; **significant at P B 0.01; be attributed to the fact that crop maturity is an
***significant at P B 0.001 agronomic trait, where a plant undergoes progressive
DAF days to 50% flowering, LS days to leaf senescence, TTW growth stages from emergence to senescence, charac-
total tuber weight, ATW average tuber weight terized by their reproductive capacity and phenology
(Struik, 2010; Khan et al. 2013). This could also be
due to the differences in environmental conditions as
of additive genetic effects observed for these traits has plant senescence takes place faster at higher temper-
been reported in previous studies (Killick 1977; Gopal atures (Kooman and Haverkort 1995).
1998; Hirut 2015; Muhinyuza et al. 2016). These
studies reported GCA to be more important in
magnitude than SCA in affecting potato yield. How- Conclusion
ever, some authors found both GCA and SCA to be
significant for potato yield with GCA being less The significant differences observed among general
important in magnitude than SCA (Bradshaw and combining ability (GCAf, and GCAm), and specific
Mackay 1994; Ortiz and Golmirzaie 2004; Ruiz de combining ability (SCA) effects for the genotypes
Galarreta et al. 2006; Haydar et al. 2009). In other points to the presence of sufficient genetic variation,
studies, significant SCA effects for yield have been which can be exploited for crop improvement. Both
reported (Gopal 1998; Ruiz de Galarreta et al. 2006; additive genetic effects were important in inheritance
Muthoni et al. 2015). Differences between progenies of the traits measured. For traits where additive
for tuber yields and number of tubers per plant were genetic effects were predominant, improvement can
found to be dominated by SCA effects, while for be made by selection and traits transferred to the
average tuber weight and specific gravity the GCA respective progenies. For characters that were largely
effect was more important (Tai 1976). The variations influenced by SCA and non-additive genetic action,
in the significance of GCA and SCA observed in further genetic gains can be achieved through
several studies might be due to differences in genetic hybridization of the desirable parents. Parents
material used (Neele et al. 1991; Ortiz and Golmirzaie, 393,077.54, 396,038.107, Rwangume and NKRK
2004; Muthoni et al. 2015). The broad sense heritabil- 19.17 had desirable GCA effects for days to flowering,
ity estimates revealed number of days to 50% flow- total and average tuber weight, indicating that these
ering (0.78) and total tuber weight (0.70) to be highly had desirable attributes for high yield and early
heritable traits. The narrow sense heritability estimates maturity. Families of Rwangume 9 NKRK19.17,
obtained in this study were varying and relatively low. 393,077.54 9 395,011.2, 396,038.107 9 Rwangume
This could be due to the genetic differences among the and 396,038.107 9 395,011.2 had desirable SCA
materials used and the environmental variations. effects for yield and number of days to 50% flowering.
Several authors (Ortiz et al. 1997; Bradshaw et al. The selected parents and families will be subjected to
2000; Iragaba, 2013) found comparatively higher further clonal evaluation and selection.
values for leaf senescence, days to flowering, total
Acknowledgements The Alliance for a Green Revolution in
tuber yield, marketable and average tuber yield among Africa (AGRA) is sincerely thanked for funding this study. Due
different populations. thanks to the National Agricultural Research Organization
The significant positive correlations between yield (NARO) for all the support.
related traits observed have been reported in other

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nitrogen and FYM interaction on yield and yield traits of
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