Journal of Research in Ecology
An International Scientific Research Journal
Variation analysis of phenological traits in the potatoes of Iran
Authors:
Journal of Research in Ecology
Bahram Dehdar1,
Jaber Panahandeh2 and
Alireza Motallebi Azar2
Institution:
1. Ardebil Agricultural and
Natural Resources Center,
Agricultural Research,
Education and Extension
Organization (AREEO),
Ardebil, Iran.
2. Department of
Horticulture,
Faculty of Agriculture,
University of Tabriz,
Islamic Republic of Iran.
Corresponding author:
Bahram Dehdar
Email ID:
Web Address:
http://ecologyresearch.info/
documents/EC0378.pdf
Journal of Research
in Ecology
An International
Scientific Research Journal
ISSN No: Print: 2319 –1546; Online: 2319– 1554
Original Research
ABSTRACT:
The potato production requires earliness and late maturity cultivars,
depending on the type of culture and environment. Hence this research was made to
find out the genotypic and phenotypic variances, heritabilities and genetic advance
for phenological traits. In the following, General Combining Ability (GCA) of parents
and Specific Combining Ability (SCA) from each crosses was calculated. Moderate and
low phenotypic coefficient of variation was observed for all of the traits except the
flowering period. The heritability estimates were found higher for all the characters
studied and varied from 77% to 58% indicating that the characters are less influenced
by environmental factors. The AS692, AS72,HS and Stbr2 cultivars may be good
combiners for earliness and in return, Agria, UT43 and UT42 may be good combiners
for late maturity breeding programs. Agria×UT42, AS72×Caeser, Daifela×Stbr2,
Luca×UT43, Satina×AS12, Satina×AS14, Satina×AS692 and Satina×Stbr2 were the best
crosses because of the most negative SCA effects of the studied traits.
Keywords:
GCA, potato, physiological traits, SCA.
Article Citation:
Bahram Dehdar, Jaber Panahandeh and Alireza Motallebi Azar
Variation analysis of phenological traits in the potatoes of Iran
Journal of Research in Ecology (2017) 5(2): 1156-1165
Dates:
Received: 26 May 2017 Accepted: 15 July 2017 Published: 01 Oct 2017
This article is governed by the Creative Commons Attribution License (http://creativecommons.org/
licenses/by/4.0), which gives permission for unrestricted use, non-commercial, distribution and
reproduction in all medium, provided the original work is properly cited.
1156-1165| JRE | 2017 | Vol 5 | No 2
www.ecologyresearch.info
Dehdar et al., 2017
INTRODUCTION
Preliminary crossing of parents possessing com-
plementary traits based on the phenotype followed by
selection in subsequent clonal generations is one of the
potato breeding methods (Sleper and Poehlman, 2006).
Identifying of clones for release as new varieties of po-
tato, can be done with selection is based on phenotype
values (Lynch and Walsh, 1998; Bradshaw and
Bonierbale, 2010). Since the selection of parents are
pertained on their performance, it is difficult to prognos-
ticate the segregation template of the F1 progeny
because the potato is a highly heterozygous crop
(Wolfgang et al., 2009). Successful breeding is attained
through incorporating the eligible alleles into a single
genotype and testing their stability and adaptation
(Acquuah, 2007). Dominance and epistatic effects con-
tribute to clone performance, therefore potatoes are
highly heterozygous. However, the value of the cross
should not be assumed unless the progeny has been test-
ed (Muthoni et al., 2012). Breeding of potato has been
used to develop genotypes for persistence to biotic and
abiotic stresses, earliness and high yield (Razukas and
Jundalas, 2006; Azhar et al., 2007). Among these attrib-
utes, earliness is important in areas with land paucity,
multiple cropping systems or short rainy concise. Such
genotypes would be good for highlands in Iran and else-
where where genotypes would need to mature quickly
making land readily accessible for other crops for the
next cropping season thus increasing agricultural
productivity. Earliness in shorter rainy seasons simpli-
fies drought escape, a trait that is decisive in climate
change conditions of diminished or disordered rainfall
templates (Banziger et al., 2000). Breeding for earli-
ness has also been considered advantageous, so its guid-
ance to release from some diseases that become visible
late in the season such as late blight (Razukas and
Jundalas, 2006). However, in potato breeding for earli-
ness has not been deliberately followed in Iran for rea-
sons of producing versatile genotypes mighty of stabil-
1157
ity to different environments. Attaining this needs un-
derstanding the method of heredity conditioning of this
Attribute (El-Bramawy and Shaban, 2007). It will be
great to define if the gene action for earliness is due to
additive, non-additive, dominant or epistatic effects and
their interaction with the environment. To evaluate the
efficiency of a given parent in hybrid combinations,
combining abilities (General Combining Ability (GCA)
and Specific Combining Ability (SCA)) are used in
plant breeding (Haydar et al., 2009). In a hybrid combi-
nation, GCA determines the average efficiency of a pa-
rental line and as the same way, SCA is the portion of
an inbred line to hybrid efficiency in a cross with a de-
termined inbred line in relation to its portions in crosses
with a domain of distinctive inbred lines (Sleper and
Poehlman, 2006; Panhwar et al., 2008). SCA dedicate
conditions were specified crosses do better or worse
than anticipated based on the efficiency of the parent
captive (Panhwar et al., 2008). SCA is due to non-
additive gene effects while GCA is mainly imputed to
additive gene action (Shattuck et al., 1993). In potato
breeding, both general and specified combining abilities
are fundamental in conditioning traits and they are both
constants in the F1 generation because there is no subse-
quent segregation with clonal breeding supplies. For
tuber yield and quality attributes in crosses between non
- related parents GCA seems to be significantly larger
than SCA whereas SCA appears to be more substantial
among related parents (Ortiz and Golmizaie, 2004). In
regard to crop maturity, GCA effects regard to be more
significant than SCA effects (Johansen et al., 1967).
Moreover, additive and non additive effects manage the
total yields of tuber and puberty (Buso et al., 2006).
This study was to determine the combining ability ef-
fects for phenological traits of selected potato clones
and their crosses. Selected parental clones and promis-
ing families will be used for further breeding in Iran and
similar agro-ecologies.
Journal of Research in Ecology (2017) 5(2): 1156–1165

MATERIALS AND METHODS
Seven high-yielding commercial cultivars of
potato (Agria, Savalan, Picasso, Caesar, Daifla, Satina
and Luca) and eleven progressive potato clones, (UT42
and UT43 which were obtained from the cross of
S. tuberosum L. ssp. andigena; three advanced and high
-yielding clones of S. tuberosum designated AS10,
AS12, AS14, AS20, AS72 and AS692, and HS, Stbrkz
and Stbr2 were obtained from the cross of S. tu-
berosum × S. Stoloniferum) were crossed reciprocally,
in the Ardabil Natural Resources and Agricultural Re-
search Station (located in 47° and 59‘ North, 39° and
22’ East, 1390 m above sea level) during the summer of
2012 and the best cultivars and clones were determined
when they were selected as the female or male parent.
By pushing petals aside and removing anthers,
the flower buds of parent were castrated. The castrated
flower buds placed in bags. The next day, flower buds
were pollinated and labeled. The ripe and fallen berries
were collected after about six weeks. The ripened ber-
ries were processed by cutting them and emptying the
seeds into a basin containing clean water. The harvested
seeds were washed and spread on filter papers and
placed to air-dry overnight. To break dormancy, the
seeds were soaked in 1500 ppm GA3 solution for 24
hours and thereafter they were rinsed and immediately
sown (Jackson and Hanneman, 1999). Weathering was
done and the seedlings were sprayed against pests and
diseases as required and seedlings were transplanted to
the field four weeks later. In the next year, to appraise
the genetic parameters (GCA, SCA and H2b), 30 seed-
lings from each family (54 cross) and 18 parents
(control) were planted in the augmented design with
5fivereplications. The distance between seedlings lines
were 75 cm. Agricultural operations include weeding,
ridging and pest control were done as per recommenda-
tions for potato production in Iran.
Estimation error and effect of block were evalu-
ated using control observations. Observations were rec-
Journal of Research in Ecology (2017) 5(2): 1156–1165
Dehdar et al., 2017
orded for 50% flowering, days to stolen appearance,
growing period, flowering period and number of days to
tuber formation for five random plant basis in each rep-
lication.
Variance analysis of controls and varieties were
performed on the basis of complete block design using
SPSS 22 software (SPSS, Inc., Chicago). The effect of
each block for each treatment was calculated and each
treatment was corrected based on the block effect. The
analysis of design allowed us to estimate the genetic
variances of the response variables.
The phenotypic and genotypic variances of each
trait were estimated from the RCBD analysis of vari-
ance. The expected mean squares under the assumption
of a random effects model was computed from linear
combinations of the main squares and the phenotypic
and genotypic coefficient of variations were computed
as per the methods suggested (Burton and Devane,
1953).
where, MSg = mean square due to genotype; MSe =
environmental variance (error mean square) and r =
number of replication.
Broad sense heritability was estimated based on the
formula given by (Allard 1960) as follows:
Genetic advance as a percent of the mines were estimat-
ed as descriptively by (Allard,1960) as:
where, K= the standardized selection differential of 5 %
(2.063); VP = phenotypic standard deviation and
H2B=broad sense heritability.
1158
Dehdar et al., 2017
Phenotypic variance was relatively greater than the gen-
otypic variance in magnitude for all characters consid-
th
The GCA and SCA effects of ijk observation ered. Phenotypic coefficient of variation ranged from
were estimated by adopting the following model. 4.47% for growing period to 23.69% for flowering peri-
Xijk = μ + gi + gj + sij od. The phenotypic and genotypic coefficient of varia-
where, μ= population mean; gi= GCA effect of ith fe- tion values can be categorized as low (<10%), moderate
th
male parent; gj= GCA effect of j male; sij= SCA effect (10-20%), and high (>20%) as indicated by Robinson
th th
of hybrid of i female with j male. and Barry (1966). Moderate and low phenotypic coeffi-
The individual effects of GCA and SCA were cient of variation were observed for all of the traits
estimated from the data obtained from two way tables of except of flowering period. The traits which exhibited
female vs. male as given below. In this each data was high estimates of genotypic and phenotypic coefficient
totaled over replications. of variations had a high probability of improvement
Two way table for female vs. male: through selection while traits with low estimates the

Male 1 2 3 ............... n Total improvement through selection is difficult or virtually

Female impractical due to the masking effect of environment on

1 Xij . . ............... . Xi.. the genotypic effect (Singh,1990).
2 . . . ............... . .
3 . . . ............... . . The heritability estimates were found higher for
. . . . ............... . . all the characters studied from 77% to 58% indicating
. . . . ............... . .
n . . . ............... . . that the characters are less influenced by environmental
Total X.j. . . ............... . X…
factors. The flowering period and growing period record-
ed highest and lowest genetic advance as a percentage of
the mean (37.64 and 5.63 respectively). Genetic advance
where, X… = Total of all hybrid combination; Xi.. = To- in general was low for all of the characters studied,
th
tal of i female over ‘t’ male and ‘r’ replications; X. j. = which showed a moderate genetic advance as a percent-
Total of jth male over ‘l’ female and ‘r’ replication and age of the mean. The low value of genetic advance for
th th
Xij. =Total of the hybrid between i female and j male these traits showed that these characters are not governed
over ‘r’ replications (Choudahry, 2002) by additive genes and selection could not be rewarded for
the improvement of these traits. In general, it has been
RESULTS accepted that the genetic coefficient of variation was not
Analysis of variance sufficient to test heritable variation, so more reliable pro-
Significant differences among parents for all cedures need to estimate the heritability and the genetic
studied traits were observed with analysis of variance advance. GCV, H2B and GA estimates would be effective
(Table 1). Finding of significant difference between of during the use of election in the breeding programs. In
parents indicated the presence of genetic variation making for these decisions, H2B estimated by the variance
among them that could be used in selection for favora- components method would be effective. Mishra et al.
ble traits. (2006) also derived that high GA would be valuable in
Genotypic and phenotypic coefficient of variation selection schedules. H2B estimates based on the growing
Estimates of the genetic parameters (VG, VP, genotypes at several environments (locations and years)
2
H b, GCV, PCV, GA and GAM) are shown in Table 2. will help to breeders in deciding about the breeding pro-

1159 Journal of Research in Ecology (2017) 5(2): 1156–1165

 Dehdar et al., 2017

Table 1. Mean squares for the 18 parents

Df 50% F DSA GP FR FP NDTF
Replication 4 69.66 12.57 46.84 1.56 24.23 15.29
** ** ** ** **
Cultivars 17 349.7 68.06 163.9 4.09 283.6 49.07**
Error 68 77.24 16.10 68.6 0.991 64.32 16.89
CV% 16 8 7 14 25 7
50% F: 50% Flowering, DSA: Days to stolen appearance, GR: Growing period, FR: Flowering rate,
FP: flowering period, NDTF: Number of days to tuber formation **:Significant at 1%

grams. Due to their stabile genetic structure in clones of
potato, additive, dominant and epistatic type of genetic
variability could be observed. For these reason, Ozturk
and Yildirim, (2014) Concluded that estimation of broad
sence heritability by using the variance components
method might be dependable in potatoes in comparison
with the generatively producing crops .
GCA Effects and mean performance of the parents
GCA and SCA effects of the selected parents as
combined with other genotypes were estimated in this
study. Estimates of GCA effects varied from 14.8, 11.9,
10.3 and 10.3 (Caesar F, HS F, HS M and Agria F
respectively) to -10.2, -7.53, -7.45, -6.2, -6.03 (AS72 F,
Savalan M, AS14 M, AS14 F and Satina M) to50%
flowering (Table 3). For days to stolen appearance,
GCA effects varied from 7.32, 6.57, 6.57 and 5.74
(UT43 M, Agria F, UT 43 F and UT42 M) to -4.76, -
4.63, -4.43 -4.03 (Stbr2 F, HS M, As692 and Stbr2 M).
For growing period GCA effects varied from12.99,
10.99 and 8.66 (UT43 M, F and UT42 M) to -8.51, -
7.01, -7.81 and -8.01 (AS692 M and F, Stbr2 M and F).
For flowering rate GCA effects varied from 0.92 and
0.66 (UT43 F and AS10 F) to -0.83, -0.73and -0.68
(AS72 F, AS692 M andAS12 F). For Flowering period
GCA effects varied from 8.5 and 6.5 (Stbrkz F and
UT43 F) to -9.5 and -7 (Agria F and AS692 M).Finally
for number of days to tuber formation GCA effects var-
ied from 7.44,6.94, 6.44 and 6.27 (Agria F, UT43 M,
UT43 F and UT42 M) to -5.16, -4.56 and -4.23 (Stbr2
M, AS692 M and Stbr2 F). Based on these estimates, it
seems that, AS692 and Stbr2 cultivars with the least
GCA value (in most of the traits) were considered to be
an appropriate parent for hybridization as a negative
combiner to reduce this traits. The highest GCA values

Table 2. Estimate of genotypic and phenotypic variance, heritability of 11 evaluated variables in 18

potato parents

S.No. 50% F DSA GP FR FP NDTF

1 VG 54.496 10.386 19.062 0.6198 43.86 6.436
2 VP 69.94 13.606 32.78 0.818 56.72 9.814
2
3 H B 0.78 0.76 0.58 0.76 0.77 0.76
4 GCV 13.48 6.54 3.59 11.23 20.84 4.47
5 PCV 15.27 7.48 4.71 12.9 23.69 5.52
6 GA (5%) 13.46 5.78 6.85 1.42 11.96 4.91
7 GAM% 24.57 11.74 5.63 20.23 37.64 8.66
50% F: 50% Flowering, DSA: Days to stolen appearance, GR: Growing period, FR: Flowering rate, FP: flower-
ing period, NDTF: Number of days to tuber formation
VG: Genotypic variance; VP: Phenotypic variance; H2B: Broad Since heritability; GCV: Genotypic coefficient
variation; PCV: Phenotypic coefficient variation; GA: Genetic advance; GAM: Genetic advance as percent of
mean.

Journal of Research in Ecology (2017) 5(2): 1156–1165 1160

 Table 3. Estimates of mean and general combing ability of 11 evaluated variables in 18 potato parents (female and male)

1161
Parent 50% F DSA GP FR FP NDTF
GCA Mean GCA Mean GCA Mean GCA Mean GCA Mean GCA Mean

Agria F 10.3 58.8±2.2 6.57 47±2.4 5.99 120.8±2.1 -0.38 5.84±0.6 -9.5 26±2.2 7.44 57.4±4.2
Luca F -1.2 51±4.8 -2.31 45±3 -0.01 120.4±3.2 -0.05 6.78±0.1 -5.25 27.2±1.8 -1.18 53.8±3.9
Dehdar et al., 2017

M 1.52 61±5.1 0.12 52±1.5 -3.79 118.2±4.1 -0.64 6.04±0.2 -1.17 34.4±1.7 -0.34 59±2.8
Caesar
F 14.8 61±5.3 4.07 52±2.1 2.99 118.2±0.9 0.07 6.04±0.4 1.5 34.4±2.2 3.44 59±4.1
M -6.03 50±3.1 1.24 49±4.3 1.32 122±0.6 -0.05 6.64±0.2 3.5 38.2± 0.77 57.2±3.2
Satina
F -0.87 50±1.7 0.4 49±2.7 -0.18 122±1.9 -0.15 6.64±0.6 0 38.2±2.3 0.11 57.2±1.6
M -2.2 54.6±1.1 -1.93 48±2.2 -6.01 114.2±4.2 0.09 6.94±0.8 -1 35.4±1.8 -2.31 55±3.8
Picasso
F 3.55 54.6±4.2 0.57 48±3.1 -3.76 114.2±4.8 -0.08 6.94±1 1 35.4±1.5 -0.81 55±2.6
M -7.53 48.2±1.7 -2.26 47±2.4 1.99 128±1.3 0.33 7.96±0.2 4.5 42.8±1.6 -1.06 57.8±3
Savalan
F 0.9 48.2±4.9 1.97 47±1.8 5.39 128±1.8 0.6 7.96±0.7 3.9 42.8±1.5 1.44 57.8±1.8
M 2.3 56.4±3.7 0.9 46±0.9 4.66 125±3.2 0.62 7.76±0.6 3.5 38.8±3.2 1.11 55.6±1.5
Daifla
F 1.3 56.4±1.2 -2.18 46±4.1 0.99 125±2.8 0.39 7.76±1.1 -0.5 38.8±2.3 -1.06 55.6±2.1
AS10 F -1.7 55.2±5 1.97 56.8±2.2 2.99 131.6±4.1 0.66 8.82±0.6 5.7 42.8±2.7 1.44 63.8±2.4
M -5.7 47.6±3.4 -1.43 52.2±1.7 -0.01 124.6±1.3 0.02 7.14±0.3 -2.5 32.4±2.1 -1.06 61.2±3

results of this study indicated that the AS692, AS72, HS

and containing positive alleles for them and in return,

of traits was observed in Agria, UT43 and UT42 (Table
3). These parents could be use in hybridization pro-

and Stbr2 cultivars may be good combiners for earliness

grams for improving and enhancing of these traits. The
AS20
F -3.7 47.6±2.9 1.07 52.2±1.2 -3.01 124.6±2.3 -0.63 7.14±0.4 -0.5 32.4±1.8 0.94 61.2±2.7
M -1.7 61±3.7 -1.43 54.6±4 -2.01 127.8±1.2 0.02 7.88±0.6 1.5 34.2±2.6 -1.56 61±4.2
AS12
F 6.3 61±2.8 2.57 54.6±0.4 0.01 127.8±2.5 -0.68 7.88±1.4 -0.5 34.2±2.1 1.44 61±3.7
M -5.2 75±4.3 -4.43 47.2±5.1 -8.51 117.8±0.6 -0.73 6.66±0.1 -7 24.6±2.1 -4.56 55.6±1.8
AS692
F -2.7 75±2.8 -2.76 47.2±3.1 -7.01 117.8±0.8 -0.32 6.66±0.5 -4.5 24.6±2.3 -2.56 55.6±2.6
M 11.9 51±1.2 -4.63 46.8±4.1 -3.61 113.6±2.6 0.2 6.24±0.7 -0.1 22.8±1.8 -3.16 47.2±2.4
HS
F 10.3 51±1.8 -.343 46.8±2.3 -3.51 113.6±4.1 -0.33 6.24±0.9 -1.5 22.8±2.1 -2.56 47.2±2.1
M -5.7 47.2±4.1 -1.43 52.6±2.9 -3.01 120.6±3.1 0.42 7.18±0.6 -4.5 26±2 -2.56 57.8±2.7
Stbrkz
F -5.2 47.2±2.6 0.07 52.6±3.4 -0.01 120.6±5.1 0.42 7.18±0.4 8.5 26±2.3 -0.56 57.8±3.8
M -2.3 60.8±2.5 -4.03 43.4±3.4 -7.81 111±2.3 -0.4 5.34±0.6 -3.3 16.2±1.9 -5.16 51.8±2.4
Stbr2
F 0.3 60.8±1.8 -4.76 43.4±2.8 -8.01 111±2.6 -0.58 5.34±0.7 -3.83 16.2±1.7 -4.23 51.8±1.6
M -7.45 46±2.8 -0.93 53±0.6 0.49 128.4±3.1 0.14 7.7±0.5 3 39.6±3.6 -0.81 61.2±0.8
AS14
F -6.2 46±3.2 0.32 53±±0.1 1.24 128.4±1.4 0.07 7.7±0.2 3.75 39.6±2.8 0.44 61.2±0.9
UT42 M 6.3 60.8±4.1 5.74 49.4±4.2 8.66 118.6±2.3 0.35 7.14±0.4 -2.17 24.2±2.2 6.27 56.2±2.4
M 5.55 58.8±1.8 7.32 50.4±2.2 12.99 128.2±2.5 0.52 7.9±0.5 3.25 33.6±2.3 6.94 56.2±2.6
UT43
F 0.3 58.8±0.7 6.57 50.4±3.1 10.99 128.2±3.4 0.92 7.9±0.3 6.5 33.6±1.9 6.44 56.2±2.3
AS72 F -10.2 37±2.8 -3.43 45±0.8 -0.51 121.6±1.8 -0.83 5.96±0.9 -2 27.1±1.8 -2.56 53.4±3.1
50% F: 50% Flowering, DSA: Days to stolen appearance, GR: Growing period, FR: Flowering rate, FP: flowering period, NDTF: Number of days to
tuber formation
F and M: Female and Male respectively

Journal of Research in Ecology (2017) 5(2): 1156–1165

potential parents enables the breeder recognition of bet-
maturity breeding programs according to the result of

ter parental forms, next introducing them into crossing

programs in order to generate genetic variation in new
this research. Information on combining abilities of the
Agria, UT43 and UT42 may be good combiners for late
 Dehdar et al., 2017
Table 4. Estimates of mean and specific combing ability of 11 evaluated variables in 54 potato cross
Female*male 50% F DSA GP FR FP NDTF
SCA Mean SCA Mean SCA Mean SCA Mean SCA Mean SCA Mean
Daifela×AS20 0.29 52±2.5 -0.51 50±2.1 0.08 126±5.2 0.08 7.3±0.61 0.81 34±1.1 -0.38 59±2.1
Daifela×Stbr2 -8.41 47±3.6 -1.3 45±2.4 0.48 120±4.6 -0.04 7±0.72 -1.12 30±1.3 -0.75 54±2.3
Daifela×HS -1.51 66±4.2 0.34 47±3.1 0.86 123±3.7 0.24 7.7±0.62 1.11 35±1.8 -0.58 56±1.5
Daifela×UT42 0.29 63±1.8 -1.43 55±1.7 -1.08 135±4.3 0.022 7.9±0.25 0.48 33±2.1 -0.71 65±1.8
Luca×AS14 0.41 47±4 1.28 50±1.2 0.54 126±2.8 0.022 7.1±0.46 0.69 32±2.2 -0.05 58±2.3
Luca×AS692 2.54 52±2.3 1.57 47±1.6 1.17 118±4.6 -0.04 6.4±0.75 1.81 24±2.6 0.33 55±2.5
Luca×AS20 -0.71 48±3.5 1.15 50±2.5 0.92 124±4.3 0.23 6.9±0.56 1.56 28±1 -0.17 58±0.9
Luca×StbrKz 2.04 50±3.1 1.65 50±2.1 -1.08 122±2.7 0.11 7.5±0.43 -0.94 29±2.2 0.33 57±3
Luca×Stbr2 2.59 55±2.7 0.37 45±2.3 0.32 117±4.1 0.02 6.5±0.52 -1.37 23±2.1 0.46 54±3.1
Luca×HS -0.51 64±3.1 -3.99 41±2.7 -0.03 121±3.5 0.09 7±0.28 1.86 29±1.5 -0.37 55±2.5
Luca*UT42 0.29 60±2.2 0.23 55±3.1 0.75 134±3.7 -0.02 7.3±0.54 2.23 28±1.4 0.5 65±2.7
Luca×UT43 3.66 60±3 -0.97 55±1.2 1.42 138±3.1 -0.09 7.6±0.7 0.19 33±2 0.08 65±2.2
Satina×AS12 0.54 54±1.5 -2.72 50±2.4 -1.16 123±2.1 0.51 7.1±0.81 -0.44 35±0.9 -1.8 58±2.6
Satina×AS14 4.66 49±1.8 -1.85 50±2.8 -3.03 123±2.5 -0.03 7±0.52 -0.31 38±0.8 -1.67 58±2.4
Satina×AS692 1.79 49±1.2 -1.56 47±1.3 -2.41 115±2. -0.09 6.3±0.63 -0.19 29±1.2 -1.23 55±2.5
Satina×Stbr2 5.84 56±3.1 -2.76 45±1.9 -3.26 114±2 0.07 6.5±0.29 -1.37 30±1.4 -1.16 54±2.1
Satina×UT42 3.54 61±2.7 1.11 59±2.5 1.17 135±2.6 0.13 7.4±0.53 1.23 34±0.9 -0.13 66±2.3
Satina×UT43 5.91 60±1.5 0.9 60±2.4 1.84 139±2.9 -0.34 7.3±0.46 -4.81 35±2. 0.45 67±2.3
Picasso×AS14 2.54 51±1.2 0.65 51±2.8 2.42 123±5.1 -0.03 7.1±0.49 -0.56 36±0.7 1.33 59±2.4
Picasso×Stbr2 -1.28 53±1.9 4.74 51±1.4 2.19 114±3.7 -0.14 6.4±0.63 6.38 36±0.5 0.84 54±2.8
Picasso×HS 4.61 71±2.3 1.38 48±2.6 1.84 118±4.3 0.24 7.2±0.62 0.61 33±2.1 1 56±2.6
Picasso×UT42 0.41 62±2.8 1.61 58±3.4 1.62 130±5.2 -0.08 7.3±0.51 1.98 33±1.4 1.87 66±2.5
Caeser×HS 1.13 75±3.7 0.6 50±2.7 -0.62 120±3.8 0.33 7±0.74 0.45 33±1.8 -0.11 58±1.9
Caeser×UT43 0.3 66±4.1 0.63 61±2.6 -0.19 136±3.4 -0.16 7.3±0.41 -1.23 37±2.1 0.34 68±1.8
Agria×UT42 -5.21 66±3.4 -5.64 58±2.1 -8.24 131±5. 0.26 6.7±0.29 2.48 24±2.3 -6.13 67±2
Savalan×AS14 1.53 46±2.9 0.12 51±1.6 0.84 130±5.1 0.11 7.7±0.49 -0.76 40±2.4 0.58 60±0.7
Savalan×Stbr2 5.7 56±4.2 4.21 51±1.7 0.62 121±4.9 0.01 7±0.35 -1.82 32±2.5 1.09 56±3
Savalan×HS -0.39 62±4.1 0.84 48±2.5 0.27 125±4.2 0.08 7.5±0.51 0.41 37±2.1 0.25 57±2.5
Savalan×UT42 2.4 60±3.7 1.07 58±2.9 0.05 137±3.1 0.16 8±0.9 0.78 36±2.3 0.12 66±2
Savalan×UT43 4.78 59±3.5 0.87 59±2.5 -1.28 139±3.6 -0.01 8.2±0.74 -0.26 42±1.9 -0.3 66±2.6
AS72×Savalan -26.29 43±2.9 -0.76 47±3 -0.78 127±5 0.14 6.8±0.65 -1.37 34±1.4 -0.05 57±1.8
AS72×Caeser -4.57 48±2.8 -1 49±2.1 -1.69 122±2.9 -0.2 5.7±0.25 -2.35 29±2.1 -1.41 57±2.8
AS10×Savalan 2.4 52±3.4 -0.16 53±2.7 0.72 132±3.6 -0.24 7.9±0.6 -1.09 42±1.6 0.95 62±2.6
AS10×Picasso 0.41 54±2.8 0.38 53±2.2 1.295 124±3.8 0.11 7.8±0.28 -0.89 38±1.9 0.7 60±2.4
AS10×Caeser -2.07 59±4 -0.4 55±1.3 -1.19 126±3.1 -0.09 7.3±0.48 -2.051 37±2.3 -0.41 62±2.3
AS10×Satina 1.54 51±3.1 -0.12 54±1.4 -0.16 128±2.5 0.02 7.6±0.56 -1.64 39±2.8 -0.3 61±2.5
AS10×Daifela -0.71 54±4.2 -0.66 52±5.2 -0.41 130±4.2 -0.09 8.1±0.34 -0.39 40±2.5 -0.88 60±2
AS14×Savalan 1.53 46±1.9 0.12 51±2.3 0.84 130±4 0.31 7.9±0.74 -0.76 40±2.3 0.58 60±2.3
AS14×Picasso 2.54 51±2.8 0.65 51±2.2 2.42 123±2.6 -0.13 7±0.36 -0.56 36±2.1 1.33 59±2.2
AS14×Caeser -3.94 52±2.7 -0.12 53±1.2 -0.06 125±5.1 -0.14 6.7±0.54 -1.73 35±2.6 0..22 61±3
AS14×Satina 4.66 49±2.6 0.15 52±1.8 0.96 127±4.7 -0.02 7±0.67 -0.31 38±2.8 0.33 60±1.9
AS692×Savalan 1.65 49±3.2 0.41 48±4 1.47 122±5.3 -0.06 6.9±0.63 0.36 32±1.5 0.95 57±2.4
AS692×Picasso 1.66 53±3.4 0.94 48±1.3 3.04 115±4.2 0.9 7.4±0.52 0.56 28±1.7 1.7 56±3.1
AS692×Caeser -1.82 57±1.6 0.17 50±2.6 0.56 117±4.2 -0.21 6±0.46 -0.6 27±2.5 0.59 58±2.7
AS12×Caeser -3.07 62±1.4 0.004 54±2.1 1.81 125±4 -0.01 6.4±0.47 -0.85 33±2.1 0.09 61±2.4
AS20×Caeser -3.07 55±1.7 -0.25 53±2.7 -2.69 120±1.9 -0.03 6.4±0.62 0.15 32±2 0.09 61±2.5
AS20×Satina 2.54 49±2.2 0.03 52±1 0.34 124±3.6 -0.11 6.5±0.34 0.56 34±3.1 0.2 60±2
Stbrkz×Savalan 0.15 46±2.6 -0.51 50±1.6 3.22 130±3.8 0.09 8±0.71 7.61 47±1.9 0.95 59±2.6
Stbrkz×Caeser -2.32 55±2.4 0.25 53±2.4 -2.69 120±2.5 0.155 7±0.29 1.65 37±2.5 -0.41 59±2.4
Stbr2×Savalan 2.7 53±2.8 -0.79 46±3.1 0.62 121±2.9 0.01 7±0.56 -0.82 33±2.8 1.09 56±1.5
Stbr2×Picasso 1.71 56±2.7 -0.26 46±2.3 2.19 114±4 -0.04 6.5±0.35 -1.62 28±1.9 0.84 54±1.9
Stbr2×Caeser -2.77 59±3.6 -1.03 48±2.1 -0.29 116±2.6 -0.24 6±0.49 -1.78 28±2.1 -0.27 56±1.5
HS×Caeser -5.87 68±2.8 -0.4 49±2.8 -0.64 120±4.8 -0.17 6.5±0.5 0.45 33±2 -0.11 58±2.6
HS×Daifela -3.51 64±4 0.34 47±2.4 -0.86 123±3.5 -0.46 7±0.27 -2.89 31±1.5 -0.58 56±3.2
UT43×Daifela -3.33 56±1.5 0.36 58±1.7 -3.41 136±2.6 -0.24 8±0.51 0.44 40±1.7 -0.13 66±2.1
50% F: 50% Flowering, DSA: Days to stolen appearance, GR: Growing period, 3.8 FR: Flowering rate, FP: flowering period, NDTF: Number of
days to tuber formation

breeding populations. Combining ability of a genotype
showed its capableness in cross combination with other
genotypes and helps in selection of best parents for
crossing and selecting a proper breeding program.
Sprague (1966) reported that GCA estimates the mean
performance of a parent relatively to all its hybrid com-
binations and indicates additive genetic effects (direct
measurement of the breeding value of a parent).
SCA effects and mean performance of the crosses
The specific combining ability (SCA) for study-
ing the traits for all the parental crosses was shown in
table 4. For 50% flowering the most negative SCA val-

Journal of Research in Ecology (2017) 5(2): 1156–1165 1162

Dehdar et al., 2017
ues were obtained from the crosses between
AS72×Savalan (-26.29), Daifela×Stbr2 (-8.41),
Agria×UT42 (-5.21) and HS×Caeser (-5.87). In return,
positive SCA values were obtained from the crosses
between Satina×Stbr2 (5.84), Satina×UT43 (5.91) and
Savalan×Stbr2 (5.7). For days of stolen appearance neg-
ative SCA values were obtained from the crosses be-
tween Agria×UT42 (-5.64) and Luca×HS (-3.99) and
positive SCA were obtained between Picasso×Stbr2
(4.74) and Savalan×Stbr2 (4.21).For growing period
negative SCA values obtained from the crosses between
Agria×UT42 (-8.24) and UT43×Daifela (-3.41) and as
the same way positive SCA values observed from the
crosses between AS692×Picasso (3.04) and
Stbrkz×Savalan (3.22). For flowering period negative
SCA values were obtained from the cross between Sati-
na×UT43 (-4.81) and positive SCA values were ob-
tained from the crosses between Stbrkz×Savalan (7.61)
and Picasso×Stbr2 (6.38).For number of days to tuber
formation negative SCA values were achieved from
cross between Agria×UT42 (-6.13) and positive SCA
values were obtained from crosses between Picas-
so×UT42 (1.87), AS692×Picasso (1.7), Picasso×AS14
(1.33) and AS14×Picasso (1.33). According of the re-
sults,Agria×UT42, AS72×Caeser,Daifela×Stbr2, Lu-
ca×UT43,Satina×AS12, Satina×AS14,Satina×AS692
and Satina×Stbr2 were the best crosses because of the
most negative SCA effects of the studied traits.
DISCUSSION
The heritability estimates in this research were
found moderate to high for all the characters studied
indicated that the characters are less influenced by envi-
ronmental factors. High estimates of the coefficient of
variability, heritability and genetic advance for plant
traits indicated that traits are largely controlled by addi-
tive gene action and that strength selection for them
would be effective. In return, low estimates indicate that
these traits are influenced by environmental factors and
1163
selection programs is not effective for breeding of them.
It seems that, AS692 and Stbr2 cultivars were consid-
ered to be suitable parents, with the least GCA value,
for hybridization as negative combiner to reduce the
earliness attributes. The results of this study indicated
that the AS692, AS72, HS and Stbr2 cultivars may be
good combiners for earliness and containing positive
alleles for them and in return, Agria, UT43 and UT42
may be good combiners for late maturity breeding pro-
grams according to the result of this research. Accord-
ing of the results, Agria×UT42, AS72×Caeser,
Daifela×Stbr2, Luca×UT43, Satina×AS12, Sati-
na×AS14, Satina×AS692 and Satina×Stbr2 were the
best hybrids because of the most negative SCA effects
for earliness attributes. In general, it has been admitted
that the GCV was not adequate to assay heritable varia-
tion, so more dependable methods need to estimate the
H2B and the GA. Estimates of H2B, GCV and GA
would be useful to the breeder during the application of
selection in the breeding programs. Mishra et al. (2006)
also derived that high H2B and GA would be valuable
in selection programes. Potato clones could have the
additive, dominant and epistatic type genetic variability
due to their constant genetic structure. For that reason,
estimation of heritabilities by using the variance compo-
nents method might be more reliable in potatoes in com-
parison to the generatively producing crops (Ozturk and
Yildirim, 2014).Information on combining abilities of
the potential parents enables the breeder recognition of
better parental forms, next introducing them into cross-
ing programs in order to generate genetic variation in
new breeding populations. In F1 hybrid breeding, analy-
sis of combining ability has been used in practical plant
improvement programs to determine the relative im-
portance of General Combining Ability (GCA), Specific
Combining Ability (SCA) of the parents in the perfor-
mance of F1 hybrids, and superior parents for crossing
in hybridization programs (Yoshioka et al., 2010). Gen-
eral combining ability is the manifestation of the addi-
Journal of Research in Ecology (2017) 5(2): 1156–1165

tive gene action for the selection of parents and SCA
represents the non-additive gene action (Raghvendra et
al., 2011).
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