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Int. J. Agri. Agri. R.
Muneer et al. Page 57
RESEARCH PAPER OPEN ACCESS
Line × tester analysis for yield contributing morphological
traits in Triticum aestivum under drought conditions
Muhammad Atif Muneer*1,2
, Zaib-Un-Nisa3
, Muhammad Zeeshan Munir1,4
,
Muhammad Imran1,2
, Anisa Intikhab5
, Saima Adil6
, Noor-Ul-Ain Saifullah7
1
Department of Plant Breeding and Genetics, University of Agriculture, Faisalabad, Pakistan
2
Center for Plant Biology, School of Life Sciences, Tsinghua University, Beijing, China
3
Sorghum Research Sub-station, Dera Ghazi Khan, Pakistan
4
School of Biological Sciences and Technology, Beijing Forestry University, Beijing, China
5
Department of Plant Breeding and Molecular Genetics, University of Poonch, Rawalakot, Pakistan
6
Department of Life Sciences, Islamia University, Bahawalpur, Pakistan
7
Department of Soil Sciences, Bahauddin Zakariya University, Multan, Pakistan
Article published on August 14, 2016
Key words: Line × tester analysis, Triticum aestivum, Drought, GCA, SCA
Abstract
The present study was carried out for the development of the water stress wheat cultivars with higher grain yield by studying
the genetic basis of crucial morphological traits. Nine wheat genotypes were grouped into six lines and three testers and these
parents were crossed line x tester fashion. Eighteen crosses including nine parents were planted in the field in randomized
complete block design with three replications. Three drought tolerant varieties Chakwal-50, Chakwal-86 and Kohistan-97 were
also sown to compare the results in water stress environment. Highest negative GCA effects were observed in WN-36 for plant
height (-6.17) and flag leaf area (-1.53), while for peduncle length it was noted in 8126 lines (-1.15). Highest positive GCA effects
were observed in WN-32 for a number of grains per spike (5.21), grain yield per plant (2.08) and for spikelet per spike (0.33),
while for 8126 and WN-10 the number of tillers per plant (0.67) and spike length (0.25) was found, respectively. The crosses
9451 × WN-25, WN-36 × 8126, WN-10 × 8126 showed highest negative SCA effects for plant height (-8.06), flag leaf area (-
2.89), and peduncle length (-2.05), respectively. Moreover, the cross combinations of WN-36 × WN-25, WN-32 × WN-25 and
AARI-7 × 9526 showed positive SCA effects for number of tillers per plant (1.52), spike length (0.72) and number of spikelet per
spike (0.84) respectively, while the interaction of WN-35 × 8126 crosses showed highest positive SCA effects for number of
grains/spike (5.69) and grain yield/plant (2.75). The parental material used in this study and cross combinations obtained from
these parents may be exploited in future breeding endeavors.
* Corresponding Author: Muhammad Atif Muneer  m_atifmuneer@yahoo.com
International Journal of Agronomy and Agricultural Research (IJAAR)
ISSN: 2223-7054 (Print) 2225-3610 (Online)
http://www.innspub.net
Vol. 9, No. 2, p. 57-64, 2016
Int. J. Agri. Agri. R.
Muneer et al. Page 58
Introduction
Wheat (Triticum aestivum L.) which is famously
known as the king of cereals is very important food
crop of the world’s community. It is a promising
cereal food crop with regard to production,
utilization, nutritive value, storage space qualities,
adaptation and transaction (Hogg et al., 2004)
.Wheat being stapled food is grown in every part of
Pakistan. It covers the 70% of rabi and 37% of the
total cropped area of the country (Akhtar et al., 2010).
The yield of wheat and the end-use quality is
dependent upon the genotype, environment and their
interaction(Randhawa et al., 2002).
Development of the drought resistant wheat cultivars
is a complex process when the purpose is to combine
the drought tolerance with higher grain yield.
Numerous yield contributing parameters such as
plant height, the number of tillers per plant, days to
maturity, spikelet per spike, and grains per spike and
1000-grain weight have been reported connected to
water stress tolerance of wheat (Ahmad et al., 2006).
Due to water stress the number of nodes decreases
and inter-nodal length also shorten which results in
decreased height of the main stem of the wheat plant.
It also causes the production of low dry matter due to
a decrease in plant stem height (Day and Intalap
1970). Water stress also causes the production of
desiccated grains and also results in the early
maturity of the crop (Ahmad et al., 2007).
Kempthorne described that line × tester analysis is a
very useful approach and can be utilized very
effectively (Kempthorne, 1957). Therefore, the
present study was conducted to find the gene action
for water stress environment related important
morphological traits in present genotypes/lines so
that excellent breeding strategies may be designed for
developing high yielding drought tolerant genotypes.
Material and methods
Breeding material
Six lines of wheat like AARI-7(2008-09), WN-
10(2008-09), WN-32(2008-09), WN-35(2008-09),
WN-36(2008-09) and 9451 used as female parents
and three testers e.g. 8126, 9526, and WN-25 as male
parents.
The crossing was performed and also some necessary
preventative measures were also taken at the time of
sowing to keep the genetic material free from
contamination. For the production of sufficient
hybrid seed, hand emasculation and pollination was
done.
Evaluation of F1 hybrids in the field
The hybrids seeds (F0) obtained from crosses along
with their parents were sown in the field of the
experimental area during crop season of 2012-2013
according to randomized complete block design
involving three replications. Each replication
comprised of 18 crosses, 6 parental genotypes, and
three check varieties. The plant to plant and row to
row distance was 15 and 30 centimeter, respectively.
At the time of sowing two seeds per hole were sown
and after germination thinning was done leaving one
plant at a site for good plant health and for the
purpose of drought only first irrigation was applied
after 25 days of sowing and on succeeding stages no
water was applied.
After the crop attained maturity, five guarded plants
from each row were selected and data were recorded
at an appropriate time for yield related morphological
traits. Some morphological traits like Plant Height
(cm), number of tillers per plant, peduncle length
(cm), flag leaf area (cm2), spike length (cm), number
of spikelet per spike, number of grains per spike and
grain yield per plant (g) were recorded.
Statistical analysis
The data thus collected were subjected to analysis of
variance technique as given by Steel et al. (1997), for
all the characters studied to observe the significant
differences among crosses and parents. Combining
ability estimates were made by using line x tester
analysis as outlined by (Kempthorne, 1957).
Results and discussion
A review of Table 1 shows that female parents (lines)
showed highly significant results for plant height, flag
leaf area, number of tillers per plant, number of
spikelet per spike, number of grains per spike,
Int. J. Agri. Agri. R.
Muneer et al. Page 59
grain plant height, number of tillers per plant, flag
leaf area, grain yield per spike and peduncle length
showed significant results. The interaction of line x
tester showed significant results for plant height,
spike length,
spikelet per spike but showed highly significant
results for a number of grains per spike and grain
yield per plant, while showed non-significant results
for a number of tillers per plant, flag leaf area and
peduncle length.
Table 1. Analysis of variance for line × tester experiment for various morphological traits in Triticum aestivum
under drought.
Crosses df Plant height
(cm)
Number of tillers
per plant
Spike length
(cm)
Flag leaf
area (cm2)
Spikelet per
spike
Number of grains
per spike
Grain yield
per plant (g)
Peduncle
Length (cm)
Replication 2 52.14N.S 1.12 N.S 0.75 N.S 4.70 N.S 0.10 N.S 19.46 N.S 2.87 N.S 2.46 N.S
Genotypes 26 120.32** 3.47* 0.87* 58.95** 1.30** 94** 14.37** 7.12**
Parents 8 150.74** 4.30* 1.01* 95.97** 1.58** 131.79** 11.92** 3.07 N.S
P. Vs C. 1 1.024N.S 1.66 N.S 0.55 N.S 41.91 N.S 0.37 N.S 3.85 N.S 43.97** 33.54**
Crosses 17 113.02** 3.18 N.S 0.82 N.S 42.53** 1.225** 81.52** 13.79** 7.47**
Lines 5 145.44* 1.24 N.S 0.28 N.S 56.66** 1.59* 99.60* 14.19* 7.10 N.S
Testers 2 164.19** 6.41** 0.58 N.S 152.29** 0.93 N.S 202.55 N.S 10.70* 18.09**
L x T 10 86.58* 3.51 N.S 1.13* 13.52 N.S 0.09* 48.27** 14.20** 5.53 N.S
Error 52 38.2 1.78 0.46 10.51 0.44 25.07 4.1 3.01
**Highly Significant; *Significant; N.S= Non-significant.
Table 2. Mean values of parents and crosses for various morphological traits in Triticum aestivum under
drought conditions.
Parents Flag leaf
area
Grains per spike Grain
yield/plant
Peduncle length Plant height Spike length Spikelets per
spike
Number of tillers
per plant
Females
AARI-7 21.81 55.47 12.33 12.19 90.63 11.69 18.87 8.2
WN-32 27.37 51.4 12.4 12.09 74.53 11.14 17.47 9.067
WN-35 23.79 51.13 13.13 13.27 93.87 11.19 17.2 8.667
WN-36 27.02 47.33 12.13 12.65 82.43 11.33 17.67 7.867
WN-10 21.05 38.73 12.87 12.67 89.87 11.34 17.47 8.933
9451 20.23 49.4 11.73 13.41 88.9 10.43 18.2 7.2
Males
8126 19.49 45 8.8 10.36 73.9 11.29 18.33 7.533
9526 37.4 51.6 9.6 12.27 85.53 10.2 18.4 5.333
WN-25 28.52 62.53 15.67 10.87 88.8 12.12 19.4 9
Crosses
AARI-7 × 8126 22.08 48.93 10.33 10.88 83.4 11.56 18.07 8.933
WN-32 × 8126 24.25 51.53 11.13 9.7 89.2 11.68 18.33 8.933
WN-35 × 8126 22.47 50.6 11.67 9.42 80.2 11.96 19 8.467
WN-36 × 8126 25 44 7.867 9.713 80 10.73 17.53 6.267
WN-10 × 8126 27.14 48.93 8.467 6.547 74 11.98 18.33 8.667
9451 × 8126 19.18 49.2 9.6 11.78 86.27 11.51 18.2 8.8
AARI-7 × 9526 24.73 43.2 9.533 12.43 90 11.97 19.2 7.467
WN-32 × 9526 28.62 52.13 11 11.65 87.4 10.19 17.93 5.933
WN-35 × 9526 27.36 36.73 5.8 11.13 90.13 11.09 17.13 7.4
WN-36 × 9526 33.43 48.47 10.73 9.007 75.57 10.93 17.33 6.533
WN-10 × 9526 28.53 52.33 13 12.03 91.43 11.67 18.67 8.733
9451 × 9526 27.41 49.47 11.93 12.25 86.76 11.47 17.93 7.133
AARI-7×WN-25 26.62 49.73 10.87 12.53 92.5 10.81 18.47 6.2
WN-32×WN-25 27.45 61.47 15.67 12.4 94.87 11.93 19.4 8.6
WN-35×WN-25 25.74 50.27 11.27 12.31 92.1 10.72 19 7.267
WN-36×WN-25 35.32 55.2 8.333 10.58 82.77 11.77 17.53 8.4
WN-10×WN-25 27 55.2 11.33 10.7 86.93 11.22 18.33 7.2
9451 × WN-25 28.47 49.47 10.67 9.907 77.7 11.43 18.2 7.2
The male (testers) and female (lines) parents used in
this study provided a broad range of expression
pattern for various characters as shown in Table 2.
Highest plant height was observed for WN-35(93.87
cm) while minimum plant height was shown by line
8126 (74.53 cm).
For plant height, the performance of drought tolerant
variety Chakwal-86 was best (77.97 cm) because
minimum plant height is desired due to the expected
lodging losses. Likewise, the tester 8126 (73.90 cm)
can be evaluated for future breeding programs for
developing drought tolerant genotypes with reduced
plant height.
Int. J. Agri. Agri. R.
Muneer et al. Page 60
Similarly, minimum flag leaf area is desirable for
drought tolerance due to the reason of reduced
transpiration losses from reduced area exposed to
sunlight. The female parent (line) 9451 showed the
minimum value (20.23 cm2) for flag leaf area,
followed by WN-10 and AARI-7 which is less than the
area exhibited by Chakwal-50, Chakwal-86, and
Kohistan-97.
WN-32 showed the maximum value (9.067 cm2) for
the number of tillers per plant followed by WN-25
and WN-10 which is very close to Control varieties
like Chakwal-50, Chakwal-86, and Kohistan-97.
Hence, it can be concluded that for yield purpose a
maximum number of tillers can be attained by WN-32
is recommended for drought tolerance with special
reference to this trait.
Table 3. Estimates of GCA for various morphological traits in Triticum aestivum under drought conditions.
Parents Plant height
(cm)
No. of tillers per
plant
Spike length(cm) Flag Leaf
Area(cm2)
Spikelets per
spike
Number of grains
per spike
Grain yield per
plant (g)
Peduncle
length
AARI-7 3 -0.14 0.08 -0.27 0.32 -2.53 -0.3 1.11
WN-32 4.86 0.15 -0.1 2.08 0.33 5.21 2.08 0.41
WN-35 1.85 0.03 -0.11 -0.93 0.12 -3.21 -0.9 0.12
WN-36 -6.17 -0.6 -0.22 -1.53 -0.78 -0.6 -1.5 -0.64
WN-10 -1.5 0.52 0.25 0.42 0.18 2.32 0.42 -1.07
9451 -2.04 0.03 0.1 0.22 -0.14 -0.44 0.22 0.48
8126 -3.44 0.67 0.2 -0.66 -0.01 -0.95 -0.7 -1.15
9526 1.25 -0.47 -0.14 -0.17 -0.22 -2.77 -0.2 0.58
WN25 2.18 -0.19 -0.05 0.84 0.23 3.72 0.84 0.57
Peduncle length was less than all other parents in
8126 (10.36 cm) whereas among drought tolerant
varieties Chakwal-86 showed 7.313 cm which is highly
desirable for drought tolerance. WN-25 showed the
maximum spike length, the number of spikelet per
spike, the number of grains per spike and grain yield
per plant.
Mean performance of the crosses
A review of Table 2 reveals that a considerable
variation in hybrid vigor is present in most of the
crosses for most of the characters studied. Some
hybrids/crosses even exceeded their better parents.
However, some intermediate hybrids were also
observed. Range in crosses for plant height was
observed from 74 cm (WN-10 x 8126) to 94.867 cm
(WN-32 x WN-25). Minimum value for flag leaf area
was observed in the cross 9451 x 8126. The crosses
AARI-7 x 8126 and WN-32 × 8126 showed the more
number of tillers per plant than the drought tolerant
variety Chakwal-86. So, these crosses can be further
evaluated for the future breeding program. Maximum
peduncle length and no. of grains per spike were
recorded in the crosses AARI-7 × WN-25 and WN-32
x WN-25respectively. The cross WN-10 x 8126
displayed the maximum value for spike length, and
the cross WN-32 x WN-25 had the maximum number
of spikelet per spike. It was observed that the crosses
showed better performance than their parents. A
good degree of variation was recorded for grain yield
per plant. The maximum and minimum values were
obtained from the crosses WN-32 x WN-25 and WN-
36 x WN-25, respectively.
General combining ability studies
It is clear from Table 3 that in the case of plant height,
negative general combining ability effects are more
important since more emphasis is placed upon
selection for short stature segregate in segregating
population because they ultimately turn out short
stature line which is more responsive to fertilizer and
tolerant to lodging. From this point of view, WN-36
and WN-10, 9451 among female parents and 8126
among male parents were potential parents and their
values of general combining ability were -6.17, -1.50, -
2.04 and -3.44 respectively. These results confirm the
findings of (Singh et al., 2003; Ivanovic, 2003). For
flag leaf area, negative general combining ability
effects are more important because flag leaf area is
much influenced by transpiration losses due to
exposure to sunlight and ultimately affects the grain
yield which is our main objective.
Int. J. Agri. Agri. R.
Muneer et al. Page 61
So, more emphasis is placed on the selection of
genotypes with smaller flag leaf area. From this point
of view, among the female parents, AARI-7, WN-35,
WN-36 and male parents 8128 and 9526 showed
better performance. These findings were in
accordance with the results of (Saeed et al., 2001;
Arshad and Chowdhry, 2002; Chowdhary et al.,
2007). A number of tillers per plant also plays an
important role in the grain yield as more number of
tillers are expected to result in better yielding ability.
General combining ability effects calculated for this
trait were of moderate magnitude. Among female
parents, WN-10, WN-32 and male parent 8126
exhibited higher positive general combining ability
effects. The findings of (Iqbal 2007; Srivastava et al.,
1981) supported the results reported in this write-up.
Likewise in plant height, shorter peduncle length is
preferred because an increase in peduncle length
ultimately increases the plant height and we prefer a
plant with short stature. In this study, two female
parents like WN-36 and WN-10 showed the negative
general combining ability. Likewise, one male parent
e.g. 8126 showed superior general combining ability
for this trait. So, it can be concluded that above-
mentioned parents are desirable for use in the
breeding program.
Table 4. Estimates of SCA for various morphological traits in Triticum aestivum under drought conditions.
CROSSES Plant Height
(cm)
No. of
Tillers/plant
spike length Flag leaf area
(cm2)
No. of
spikelets/spike
No of
grains/spike
Grain
yield/plant (g)
Peduncle length
(cm)
AARI-7 × 8126 -1.78 0.72 -0.09 0.96 -0.5 2.6 0.75 0.09
WN-32 × 8126 2.15 0.44 0.21 0.83 -0.21 -2.55 -0.8 -0.39
WN-35 × 8126 -3.83 0.08 0.5 0.63 0.63 5.69 2.75 -0.37
WN-36 × 8126 4 -1.47 -0.61 -2.89 0.07 -4.26 -0.44 1.1
WN-10 × 8126 -6.67 -0.2 0.15 2.94 -0.1 -2.26 -1.8 -2.05
9451 × 8126 6.13 0.41 -0.16 -2.48 0.1 0.78 -0.46 1.62
AARI-7 × 9526 0.1 0.4 0.67 -1.38 0.84 -1.31 -0.53 -0.1
WN-32×9526 -4.34 -1.41 -0.93 0.21 -0.4 -0.14 -1.4 -0.18
WN-35×9526 1.39 0.16 -0.01 0.53 -1.02 -6.36 -3.6 -0.41
WN-36×9526 -5.13 -0.05 -0.06 0.54 0.08 2.01 1.93 -1.34
WN-10×9526 6.05 1.007 0.19 -0.66 0.44 2.94 2.24 1.69
9451×9526 1.92 -0.13 0.14 0.75 0.04 2.85 1.37 0.35
AARI-7×WN-25 1.67 -1.13 -0.58 0.41 -0.34 -1.28 -0.22 0.01
WN-32×WN-25 2.19 0.97 0.72 -1.04 0.61 2.69 2.22 0.57
WN-35×WN-25 2.43 -0.24 -0.48 -1.17 0.38 0.67 0.84 0.78
WN-36×WN-25 1.13 1.52 0.68 2.35 -0.16 2.24 -1.48 0.23
WN-10×WN-25 0.62 -0.83 -0.35 -2.27 -0.34 -0.68 -0.44 0.36
9451×WN-25 -8.06 -0.31 0.01 1.72 -0.14 -3.64 -0.91 -1.97
The consequences are in line with the findings of
(Sharma and Garg, 2005). Spike length is an
important component since greater spike length has
more number of spikelet per spike and grains per
spike which ultimately results in the better grain yield
potential. Therefore, increased spike length is the
objective of a breeder. For this trait, parents with
positive general combining ability are required.
Similar studies have also been discussed by (Tosun et
al., 1995; Malik et al., 2005; Hasnain et al., 2006).
Spikelet per spike contributes positively towards
grain yield. More the number of spikelet per spike,
greater will be the grain yield. Therefore, selection for
a higher number of spikelet per spike may ultimately
lead to the evaluation of better yielding lines. Among
the female parents AARI-7, WN-10 and 9451 showed
the positive GCA values while only one male parent
8126 was found to reveal positive general combining
ability effects. These results are in the agreement with
the results reported by (Tosun et al., 1995; Saeed et
al., 2001; Malik et al., 2005; Chowdhary et al., 2007).
A number of grains per spike are also an important
factor because as the number of grains will be more,
grain yield will also increase. Therefore, positive
general combining ability effects are more important
due to the positive contribution of grain yield. Among
female parents, WN-32 and WN-10 showed positive
and higher values of
Int. J. Agri. Agri. R.
Muneer et al. Page 62
general combining ability effects for grains per spike
while among male parents, only WN-25 showed the
positive value. It should be noted that values of the
female parent were higher than a male parent. These
results match with the findings of (Saeed et al., 2001;
Singh et al., 2003; Hassan et al., 2007; Iqbal, 2007).
For grain yield per plant, three female parents (WN-
32, WN-10 and 9451) and among the male parents
WN-25 exhibited positive general combining ability
effects. Similar results were also found (Khan and
Khan, 1999; Malik et al., 2005). The highest effects
were found in the female parent WN-32 followed by
line WN-10.
Specific combining ability studies
Only six among eighteen crosses depicted negative
specific combining ability effects for plant height
(Table 4). If the parents with short stature are ideal
one, then the crosses viz., AARI-7 x 8126, WN-35 x
8126, WN-10 x 8126, WN-32 x 9526, WN-36 × 9526
and 9451 x WN-25 may be consider well. However,
the remaining crosses showed higher SCA effects.
These results confirmed the studies of (Arshad and
Chowdhry, 2002; Ahmadi et al., 2003; Hasnain et al.,
2006; Chowdhary et al., 2007). Approximately, 45%
crosses showed the smaller values of specific
combining ability effects for flag leaf area which is
desirable. As less flag leaf area is required for drought
tolerance, the crosses WN-36 x 8126, 9451 x 8126,
AARI-7 x 9526, WN-10 x 9526, WN-32 x WN-25,
WN-35 x WN-25 and WN-10 x WN-25 may be used in
the future breeding program because they have high
SCA values contributing towards minimum flag leaf
area. Similar results have also been reported by
(Saeed et al., 2001; Arshad and Chowdhry 2002;
Nazir et al., 2005; Chowdhary et al., 2007). A number
of tillers per plant are an important contributing trait.
The crosses viz., AARI-7 x 8126, WN-32 x 8126, WN-
35 x 8126, 9451 x 8126, AARI-7 x 9526, WN-35 x
9526, WN-10 x 9526, WN-32 x WN-25 and WN-36 x
WN-25 exhibited the positive values for specific
combining ability effects. These results are in
accordance with the findings of (Hassan et al., 2007).
The negative specific combining ability effects are
needed to reduce the peduncle length.
Out of eighteen crosses, eight crosses displayed
negative SCA effects. Among these, the crosses viz.,
WN-10 x 8126, WN-36 x 9526 and 9451 x WN-25 are
the best hybrids for reduced peduncle length. Similar
results were reported by (Chowdhary et al., 2007). In
the case of spike length, positive specific combining
ability effects are desired. Best crosses are those
having positive and higher values of specific
combining ability effects. Specific combining ability
was positive in 50 % of crosses. The cross WN-32 ×
WN-25 showed the highest value followed by the
crosses WN-36 × WN-25 and WN-35 x 8126. Similar
studies have also been reported by (Hasnain et al.,
2006). For a number of spikelet per spike, positive
specific combining ability effects were displayed in 9
crosses out of 18 crosses but AARI-1 x 9526, WN-35 x
8126 and WN-35 x WN-25 performed the best and
can be recommended for the future breeding
programs. Further exploration of these crosses having
positive specific combining ability effects may lead to
the selection of lines having more number of spikelet
per spike. These results are in the conformity with
those of (Singh et al., 2003); (Mahantashivayogayya
et al., 2010). Positive specific combining ability
effects were shown by 50 % of the crosses for a
number of grains per spike. Potential crosses showing
higher values of specific combining ability effects
were WN-35 x 8126, WN-10 x 9526, 9452 x 9526,
WN-32 x WN-25. The findings of (Iqbal 2007)
supported the results reported in this write-up.
Specific combining ability effects were found much
variable among crosses for grain yield per plant. The
poorest cross with respect to specific combining
ability for grain yield per plant was WN-35 x 9526
whereas the cross appeared as a best and most
promising specific combination was WN-35 × 8126.
Positive specific combining ability effects were
displayed in 07 out of 18 crosses. Such positive effects
were impressive in crosses. Same results were also
reported by (Saeed et al., 2001; Awan et al., 2005;
Hassan et al., 2007; Iqbal 2007; Singh et al., 2007;
Akbar et al., 2009). Specific combining ability effects
were found much variable among crosses for grain
yield per plant.
Int. J. Agri. Agri. R.
Muneer et al. Page 63
The poorest cross with respect to specific combining
ability for grain yield per plant was WN-32 x WN-25
whereas the cross appeared as a best and most
promising specific combination was WN-35 × 9526.
Positive specific combining ability effects were
displayed in 10 out of 18 crosses. Such positive effects
were impressive in crosses.
Conclusion
Wheat production is mainly affected by several
environmental factors and water stress is the
major constraint in Pakistan. Therefore,
understanding the mechanism of plant responses to
water stress is an important and basic part of crops.
Thus, it is need to select such genotypes of wheat
which lead to better adaptation and higher yield
against water stress. These studies revealed that
parental genotypes like WN-10, WN-25, WN-32 and
8126 are beneficial to generate desirable
combinations and may be used in future breeding
strategies. Specific combinations like, WN-35 x 8126
and WN-10 x 9526 needs further investigations.
Taken together, the information derived from these
studies may be used to develop drought tolerant
wheat material that could give economic yield in
water stressed conditions of wheat growing areas.
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Line × tester analysis for yield contributing morphological traits in Triticum aestivum under drought conditions

  • 1. Int. J. Agri. Agri. R. Muneer et al. Page 57 RESEARCH PAPER OPEN ACCESS Line × tester analysis for yield contributing morphological traits in Triticum aestivum under drought conditions Muhammad Atif Muneer*1,2 , Zaib-Un-Nisa3 , Muhammad Zeeshan Munir1,4 , Muhammad Imran1,2 , Anisa Intikhab5 , Saima Adil6 , Noor-Ul-Ain Saifullah7 1 Department of Plant Breeding and Genetics, University of Agriculture, Faisalabad, Pakistan 2 Center for Plant Biology, School of Life Sciences, Tsinghua University, Beijing, China 3 Sorghum Research Sub-station, Dera Ghazi Khan, Pakistan 4 School of Biological Sciences and Technology, Beijing Forestry University, Beijing, China 5 Department of Plant Breeding and Molecular Genetics, University of Poonch, Rawalakot, Pakistan 6 Department of Life Sciences, Islamia University, Bahawalpur, Pakistan 7 Department of Soil Sciences, Bahauddin Zakariya University, Multan, Pakistan Article published on August 14, 2016 Key words: Line × tester analysis, Triticum aestivum, Drought, GCA, SCA Abstract The present study was carried out for the development of the water stress wheat cultivars with higher grain yield by studying the genetic basis of crucial morphological traits. Nine wheat genotypes were grouped into six lines and three testers and these parents were crossed line x tester fashion. Eighteen crosses including nine parents were planted in the field in randomized complete block design with three replications. Three drought tolerant varieties Chakwal-50, Chakwal-86 and Kohistan-97 were also sown to compare the results in water stress environment. Highest negative GCA effects were observed in WN-36 for plant height (-6.17) and flag leaf area (-1.53), while for peduncle length it was noted in 8126 lines (-1.15). Highest positive GCA effects were observed in WN-32 for a number of grains per spike (5.21), grain yield per plant (2.08) and for spikelet per spike (0.33), while for 8126 and WN-10 the number of tillers per plant (0.67) and spike length (0.25) was found, respectively. The crosses 9451 × WN-25, WN-36 × 8126, WN-10 × 8126 showed highest negative SCA effects for plant height (-8.06), flag leaf area (- 2.89), and peduncle length (-2.05), respectively. Moreover, the cross combinations of WN-36 × WN-25, WN-32 × WN-25 and AARI-7 × 9526 showed positive SCA effects for number of tillers per plant (1.52), spike length (0.72) and number of spikelet per spike (0.84) respectively, while the interaction of WN-35 × 8126 crosses showed highest positive SCA effects for number of grains/spike (5.69) and grain yield/plant (2.75). The parental material used in this study and cross combinations obtained from these parents may be exploited in future breeding endeavors. * Corresponding Author: Muhammad Atif Muneer  m_atifmuneer@yahoo.com International Journal of Agronomy and Agricultural Research (IJAAR) ISSN: 2223-7054 (Print) 2225-3610 (Online) http://www.innspub.net Vol. 9, No. 2, p. 57-64, 2016
  • 2. Int. J. Agri. Agri. R. Muneer et al. Page 58 Introduction Wheat (Triticum aestivum L.) which is famously known as the king of cereals is very important food crop of the world’s community. It is a promising cereal food crop with regard to production, utilization, nutritive value, storage space qualities, adaptation and transaction (Hogg et al., 2004) .Wheat being stapled food is grown in every part of Pakistan. It covers the 70% of rabi and 37% of the total cropped area of the country (Akhtar et al., 2010). The yield of wheat and the end-use quality is dependent upon the genotype, environment and their interaction(Randhawa et al., 2002). Development of the drought resistant wheat cultivars is a complex process when the purpose is to combine the drought tolerance with higher grain yield. Numerous yield contributing parameters such as plant height, the number of tillers per plant, days to maturity, spikelet per spike, and grains per spike and 1000-grain weight have been reported connected to water stress tolerance of wheat (Ahmad et al., 2006). Due to water stress the number of nodes decreases and inter-nodal length also shorten which results in decreased height of the main stem of the wheat plant. It also causes the production of low dry matter due to a decrease in plant stem height (Day and Intalap 1970). Water stress also causes the production of desiccated grains and also results in the early maturity of the crop (Ahmad et al., 2007). Kempthorne described that line × tester analysis is a very useful approach and can be utilized very effectively (Kempthorne, 1957). Therefore, the present study was conducted to find the gene action for water stress environment related important morphological traits in present genotypes/lines so that excellent breeding strategies may be designed for developing high yielding drought tolerant genotypes. Material and methods Breeding material Six lines of wheat like AARI-7(2008-09), WN- 10(2008-09), WN-32(2008-09), WN-35(2008-09), WN-36(2008-09) and 9451 used as female parents and three testers e.g. 8126, 9526, and WN-25 as male parents. The crossing was performed and also some necessary preventative measures were also taken at the time of sowing to keep the genetic material free from contamination. For the production of sufficient hybrid seed, hand emasculation and pollination was done. Evaluation of F1 hybrids in the field The hybrids seeds (F0) obtained from crosses along with their parents were sown in the field of the experimental area during crop season of 2012-2013 according to randomized complete block design involving three replications. Each replication comprised of 18 crosses, 6 parental genotypes, and three check varieties. The plant to plant and row to row distance was 15 and 30 centimeter, respectively. At the time of sowing two seeds per hole were sown and after germination thinning was done leaving one plant at a site for good plant health and for the purpose of drought only first irrigation was applied after 25 days of sowing and on succeeding stages no water was applied. After the crop attained maturity, five guarded plants from each row were selected and data were recorded at an appropriate time for yield related morphological traits. Some morphological traits like Plant Height (cm), number of tillers per plant, peduncle length (cm), flag leaf area (cm2), spike length (cm), number of spikelet per spike, number of grains per spike and grain yield per plant (g) were recorded. Statistical analysis The data thus collected were subjected to analysis of variance technique as given by Steel et al. (1997), for all the characters studied to observe the significant differences among crosses and parents. Combining ability estimates were made by using line x tester analysis as outlined by (Kempthorne, 1957). Results and discussion A review of Table 1 shows that female parents (lines) showed highly significant results for plant height, flag leaf area, number of tillers per plant, number of spikelet per spike, number of grains per spike,
  • 3. Int. J. Agri. Agri. R. Muneer et al. Page 59 grain plant height, number of tillers per plant, flag leaf area, grain yield per spike and peduncle length showed significant results. The interaction of line x tester showed significant results for plant height, spike length, spikelet per spike but showed highly significant results for a number of grains per spike and grain yield per plant, while showed non-significant results for a number of tillers per plant, flag leaf area and peduncle length. Table 1. Analysis of variance for line × tester experiment for various morphological traits in Triticum aestivum under drought. Crosses df Plant height (cm) Number of tillers per plant Spike length (cm) Flag leaf area (cm2) Spikelet per spike Number of grains per spike Grain yield per plant (g) Peduncle Length (cm) Replication 2 52.14N.S 1.12 N.S 0.75 N.S 4.70 N.S 0.10 N.S 19.46 N.S 2.87 N.S 2.46 N.S Genotypes 26 120.32** 3.47* 0.87* 58.95** 1.30** 94** 14.37** 7.12** Parents 8 150.74** 4.30* 1.01* 95.97** 1.58** 131.79** 11.92** 3.07 N.S P. Vs C. 1 1.024N.S 1.66 N.S 0.55 N.S 41.91 N.S 0.37 N.S 3.85 N.S 43.97** 33.54** Crosses 17 113.02** 3.18 N.S 0.82 N.S 42.53** 1.225** 81.52** 13.79** 7.47** Lines 5 145.44* 1.24 N.S 0.28 N.S 56.66** 1.59* 99.60* 14.19* 7.10 N.S Testers 2 164.19** 6.41** 0.58 N.S 152.29** 0.93 N.S 202.55 N.S 10.70* 18.09** L x T 10 86.58* 3.51 N.S 1.13* 13.52 N.S 0.09* 48.27** 14.20** 5.53 N.S Error 52 38.2 1.78 0.46 10.51 0.44 25.07 4.1 3.01 **Highly Significant; *Significant; N.S= Non-significant. Table 2. Mean values of parents and crosses for various morphological traits in Triticum aestivum under drought conditions. Parents Flag leaf area Grains per spike Grain yield/plant Peduncle length Plant height Spike length Spikelets per spike Number of tillers per plant Females AARI-7 21.81 55.47 12.33 12.19 90.63 11.69 18.87 8.2 WN-32 27.37 51.4 12.4 12.09 74.53 11.14 17.47 9.067 WN-35 23.79 51.13 13.13 13.27 93.87 11.19 17.2 8.667 WN-36 27.02 47.33 12.13 12.65 82.43 11.33 17.67 7.867 WN-10 21.05 38.73 12.87 12.67 89.87 11.34 17.47 8.933 9451 20.23 49.4 11.73 13.41 88.9 10.43 18.2 7.2 Males 8126 19.49 45 8.8 10.36 73.9 11.29 18.33 7.533 9526 37.4 51.6 9.6 12.27 85.53 10.2 18.4 5.333 WN-25 28.52 62.53 15.67 10.87 88.8 12.12 19.4 9 Crosses AARI-7 × 8126 22.08 48.93 10.33 10.88 83.4 11.56 18.07 8.933 WN-32 × 8126 24.25 51.53 11.13 9.7 89.2 11.68 18.33 8.933 WN-35 × 8126 22.47 50.6 11.67 9.42 80.2 11.96 19 8.467 WN-36 × 8126 25 44 7.867 9.713 80 10.73 17.53 6.267 WN-10 × 8126 27.14 48.93 8.467 6.547 74 11.98 18.33 8.667 9451 × 8126 19.18 49.2 9.6 11.78 86.27 11.51 18.2 8.8 AARI-7 × 9526 24.73 43.2 9.533 12.43 90 11.97 19.2 7.467 WN-32 × 9526 28.62 52.13 11 11.65 87.4 10.19 17.93 5.933 WN-35 × 9526 27.36 36.73 5.8 11.13 90.13 11.09 17.13 7.4 WN-36 × 9526 33.43 48.47 10.73 9.007 75.57 10.93 17.33 6.533 WN-10 × 9526 28.53 52.33 13 12.03 91.43 11.67 18.67 8.733 9451 × 9526 27.41 49.47 11.93 12.25 86.76 11.47 17.93 7.133 AARI-7×WN-25 26.62 49.73 10.87 12.53 92.5 10.81 18.47 6.2 WN-32×WN-25 27.45 61.47 15.67 12.4 94.87 11.93 19.4 8.6 WN-35×WN-25 25.74 50.27 11.27 12.31 92.1 10.72 19 7.267 WN-36×WN-25 35.32 55.2 8.333 10.58 82.77 11.77 17.53 8.4 WN-10×WN-25 27 55.2 11.33 10.7 86.93 11.22 18.33 7.2 9451 × WN-25 28.47 49.47 10.67 9.907 77.7 11.43 18.2 7.2 The male (testers) and female (lines) parents used in this study provided a broad range of expression pattern for various characters as shown in Table 2. Highest plant height was observed for WN-35(93.87 cm) while minimum plant height was shown by line 8126 (74.53 cm). For plant height, the performance of drought tolerant variety Chakwal-86 was best (77.97 cm) because minimum plant height is desired due to the expected lodging losses. Likewise, the tester 8126 (73.90 cm) can be evaluated for future breeding programs for developing drought tolerant genotypes with reduced plant height.
  • 4. Int. J. Agri. Agri. R. Muneer et al. Page 60 Similarly, minimum flag leaf area is desirable for drought tolerance due to the reason of reduced transpiration losses from reduced area exposed to sunlight. The female parent (line) 9451 showed the minimum value (20.23 cm2) for flag leaf area, followed by WN-10 and AARI-7 which is less than the area exhibited by Chakwal-50, Chakwal-86, and Kohistan-97. WN-32 showed the maximum value (9.067 cm2) for the number of tillers per plant followed by WN-25 and WN-10 which is very close to Control varieties like Chakwal-50, Chakwal-86, and Kohistan-97. Hence, it can be concluded that for yield purpose a maximum number of tillers can be attained by WN-32 is recommended for drought tolerance with special reference to this trait. Table 3. Estimates of GCA for various morphological traits in Triticum aestivum under drought conditions. Parents Plant height (cm) No. of tillers per plant Spike length(cm) Flag Leaf Area(cm2) Spikelets per spike Number of grains per spike Grain yield per plant (g) Peduncle length AARI-7 3 -0.14 0.08 -0.27 0.32 -2.53 -0.3 1.11 WN-32 4.86 0.15 -0.1 2.08 0.33 5.21 2.08 0.41 WN-35 1.85 0.03 -0.11 -0.93 0.12 -3.21 -0.9 0.12 WN-36 -6.17 -0.6 -0.22 -1.53 -0.78 -0.6 -1.5 -0.64 WN-10 -1.5 0.52 0.25 0.42 0.18 2.32 0.42 -1.07 9451 -2.04 0.03 0.1 0.22 -0.14 -0.44 0.22 0.48 8126 -3.44 0.67 0.2 -0.66 -0.01 -0.95 -0.7 -1.15 9526 1.25 -0.47 -0.14 -0.17 -0.22 -2.77 -0.2 0.58 WN25 2.18 -0.19 -0.05 0.84 0.23 3.72 0.84 0.57 Peduncle length was less than all other parents in 8126 (10.36 cm) whereas among drought tolerant varieties Chakwal-86 showed 7.313 cm which is highly desirable for drought tolerance. WN-25 showed the maximum spike length, the number of spikelet per spike, the number of grains per spike and grain yield per plant. Mean performance of the crosses A review of Table 2 reveals that a considerable variation in hybrid vigor is present in most of the crosses for most of the characters studied. Some hybrids/crosses even exceeded their better parents. However, some intermediate hybrids were also observed. Range in crosses for plant height was observed from 74 cm (WN-10 x 8126) to 94.867 cm (WN-32 x WN-25). Minimum value for flag leaf area was observed in the cross 9451 x 8126. The crosses AARI-7 x 8126 and WN-32 × 8126 showed the more number of tillers per plant than the drought tolerant variety Chakwal-86. So, these crosses can be further evaluated for the future breeding program. Maximum peduncle length and no. of grains per spike were recorded in the crosses AARI-7 × WN-25 and WN-32 x WN-25respectively. The cross WN-10 x 8126 displayed the maximum value for spike length, and the cross WN-32 x WN-25 had the maximum number of spikelet per spike. It was observed that the crosses showed better performance than their parents. A good degree of variation was recorded for grain yield per plant. The maximum and minimum values were obtained from the crosses WN-32 x WN-25 and WN- 36 x WN-25, respectively. General combining ability studies It is clear from Table 3 that in the case of plant height, negative general combining ability effects are more important since more emphasis is placed upon selection for short stature segregate in segregating population because they ultimately turn out short stature line which is more responsive to fertilizer and tolerant to lodging. From this point of view, WN-36 and WN-10, 9451 among female parents and 8126 among male parents were potential parents and their values of general combining ability were -6.17, -1.50, - 2.04 and -3.44 respectively. These results confirm the findings of (Singh et al., 2003; Ivanovic, 2003). For flag leaf area, negative general combining ability effects are more important because flag leaf area is much influenced by transpiration losses due to exposure to sunlight and ultimately affects the grain yield which is our main objective.
  • 5. Int. J. Agri. Agri. R. Muneer et al. Page 61 So, more emphasis is placed on the selection of genotypes with smaller flag leaf area. From this point of view, among the female parents, AARI-7, WN-35, WN-36 and male parents 8128 and 9526 showed better performance. These findings were in accordance with the results of (Saeed et al., 2001; Arshad and Chowdhry, 2002; Chowdhary et al., 2007). A number of tillers per plant also plays an important role in the grain yield as more number of tillers are expected to result in better yielding ability. General combining ability effects calculated for this trait were of moderate magnitude. Among female parents, WN-10, WN-32 and male parent 8126 exhibited higher positive general combining ability effects. The findings of (Iqbal 2007; Srivastava et al., 1981) supported the results reported in this write-up. Likewise in plant height, shorter peduncle length is preferred because an increase in peduncle length ultimately increases the plant height and we prefer a plant with short stature. In this study, two female parents like WN-36 and WN-10 showed the negative general combining ability. Likewise, one male parent e.g. 8126 showed superior general combining ability for this trait. So, it can be concluded that above- mentioned parents are desirable for use in the breeding program. Table 4. Estimates of SCA for various morphological traits in Triticum aestivum under drought conditions. CROSSES Plant Height (cm) No. of Tillers/plant spike length Flag leaf area (cm2) No. of spikelets/spike No of grains/spike Grain yield/plant (g) Peduncle length (cm) AARI-7 × 8126 -1.78 0.72 -0.09 0.96 -0.5 2.6 0.75 0.09 WN-32 × 8126 2.15 0.44 0.21 0.83 -0.21 -2.55 -0.8 -0.39 WN-35 × 8126 -3.83 0.08 0.5 0.63 0.63 5.69 2.75 -0.37 WN-36 × 8126 4 -1.47 -0.61 -2.89 0.07 -4.26 -0.44 1.1 WN-10 × 8126 -6.67 -0.2 0.15 2.94 -0.1 -2.26 -1.8 -2.05 9451 × 8126 6.13 0.41 -0.16 -2.48 0.1 0.78 -0.46 1.62 AARI-7 × 9526 0.1 0.4 0.67 -1.38 0.84 -1.31 -0.53 -0.1 WN-32×9526 -4.34 -1.41 -0.93 0.21 -0.4 -0.14 -1.4 -0.18 WN-35×9526 1.39 0.16 -0.01 0.53 -1.02 -6.36 -3.6 -0.41 WN-36×9526 -5.13 -0.05 -0.06 0.54 0.08 2.01 1.93 -1.34 WN-10×9526 6.05 1.007 0.19 -0.66 0.44 2.94 2.24 1.69 9451×9526 1.92 -0.13 0.14 0.75 0.04 2.85 1.37 0.35 AARI-7×WN-25 1.67 -1.13 -0.58 0.41 -0.34 -1.28 -0.22 0.01 WN-32×WN-25 2.19 0.97 0.72 -1.04 0.61 2.69 2.22 0.57 WN-35×WN-25 2.43 -0.24 -0.48 -1.17 0.38 0.67 0.84 0.78 WN-36×WN-25 1.13 1.52 0.68 2.35 -0.16 2.24 -1.48 0.23 WN-10×WN-25 0.62 -0.83 -0.35 -2.27 -0.34 -0.68 -0.44 0.36 9451×WN-25 -8.06 -0.31 0.01 1.72 -0.14 -3.64 -0.91 -1.97 The consequences are in line with the findings of (Sharma and Garg, 2005). Spike length is an important component since greater spike length has more number of spikelet per spike and grains per spike which ultimately results in the better grain yield potential. Therefore, increased spike length is the objective of a breeder. For this trait, parents with positive general combining ability are required. Similar studies have also been discussed by (Tosun et al., 1995; Malik et al., 2005; Hasnain et al., 2006). Spikelet per spike contributes positively towards grain yield. More the number of spikelet per spike, greater will be the grain yield. Therefore, selection for a higher number of spikelet per spike may ultimately lead to the evaluation of better yielding lines. Among the female parents AARI-7, WN-10 and 9451 showed the positive GCA values while only one male parent 8126 was found to reveal positive general combining ability effects. These results are in the agreement with the results reported by (Tosun et al., 1995; Saeed et al., 2001; Malik et al., 2005; Chowdhary et al., 2007). A number of grains per spike are also an important factor because as the number of grains will be more, grain yield will also increase. Therefore, positive general combining ability effects are more important due to the positive contribution of grain yield. Among female parents, WN-32 and WN-10 showed positive and higher values of
  • 6. Int. J. Agri. Agri. R. Muneer et al. Page 62 general combining ability effects for grains per spike while among male parents, only WN-25 showed the positive value. It should be noted that values of the female parent were higher than a male parent. These results match with the findings of (Saeed et al., 2001; Singh et al., 2003; Hassan et al., 2007; Iqbal, 2007). For grain yield per plant, three female parents (WN- 32, WN-10 and 9451) and among the male parents WN-25 exhibited positive general combining ability effects. Similar results were also found (Khan and Khan, 1999; Malik et al., 2005). The highest effects were found in the female parent WN-32 followed by line WN-10. Specific combining ability studies Only six among eighteen crosses depicted negative specific combining ability effects for plant height (Table 4). If the parents with short stature are ideal one, then the crosses viz., AARI-7 x 8126, WN-35 x 8126, WN-10 x 8126, WN-32 x 9526, WN-36 × 9526 and 9451 x WN-25 may be consider well. However, the remaining crosses showed higher SCA effects. These results confirmed the studies of (Arshad and Chowdhry, 2002; Ahmadi et al., 2003; Hasnain et al., 2006; Chowdhary et al., 2007). Approximately, 45% crosses showed the smaller values of specific combining ability effects for flag leaf area which is desirable. As less flag leaf area is required for drought tolerance, the crosses WN-36 x 8126, 9451 x 8126, AARI-7 x 9526, WN-10 x 9526, WN-32 x WN-25, WN-35 x WN-25 and WN-10 x WN-25 may be used in the future breeding program because they have high SCA values contributing towards minimum flag leaf area. Similar results have also been reported by (Saeed et al., 2001; Arshad and Chowdhry 2002; Nazir et al., 2005; Chowdhary et al., 2007). A number of tillers per plant are an important contributing trait. The crosses viz., AARI-7 x 8126, WN-32 x 8126, WN- 35 x 8126, 9451 x 8126, AARI-7 x 9526, WN-35 x 9526, WN-10 x 9526, WN-32 x WN-25 and WN-36 x WN-25 exhibited the positive values for specific combining ability effects. These results are in accordance with the findings of (Hassan et al., 2007). The negative specific combining ability effects are needed to reduce the peduncle length. Out of eighteen crosses, eight crosses displayed negative SCA effects. Among these, the crosses viz., WN-10 x 8126, WN-36 x 9526 and 9451 x WN-25 are the best hybrids for reduced peduncle length. Similar results were reported by (Chowdhary et al., 2007). In the case of spike length, positive specific combining ability effects are desired. Best crosses are those having positive and higher values of specific combining ability effects. Specific combining ability was positive in 50 % of crosses. The cross WN-32 × WN-25 showed the highest value followed by the crosses WN-36 × WN-25 and WN-35 x 8126. Similar studies have also been reported by (Hasnain et al., 2006). For a number of spikelet per spike, positive specific combining ability effects were displayed in 9 crosses out of 18 crosses but AARI-1 x 9526, WN-35 x 8126 and WN-35 x WN-25 performed the best and can be recommended for the future breeding programs. Further exploration of these crosses having positive specific combining ability effects may lead to the selection of lines having more number of spikelet per spike. These results are in the conformity with those of (Singh et al., 2003); (Mahantashivayogayya et al., 2010). Positive specific combining ability effects were shown by 50 % of the crosses for a number of grains per spike. Potential crosses showing higher values of specific combining ability effects were WN-35 x 8126, WN-10 x 9526, 9452 x 9526, WN-32 x WN-25. The findings of (Iqbal 2007) supported the results reported in this write-up. Specific combining ability effects were found much variable among crosses for grain yield per plant. The poorest cross with respect to specific combining ability for grain yield per plant was WN-35 x 9526 whereas the cross appeared as a best and most promising specific combination was WN-35 × 8126. Positive specific combining ability effects were displayed in 07 out of 18 crosses. Such positive effects were impressive in crosses. Same results were also reported by (Saeed et al., 2001; Awan et al., 2005; Hassan et al., 2007; Iqbal 2007; Singh et al., 2007; Akbar et al., 2009). Specific combining ability effects were found much variable among crosses for grain yield per plant.
  • 7. Int. J. Agri. Agri. R. Muneer et al. Page 63 The poorest cross with respect to specific combining ability for grain yield per plant was WN-32 x WN-25 whereas the cross appeared as a best and most promising specific combination was WN-35 × 9526. Positive specific combining ability effects were displayed in 10 out of 18 crosses. Such positive effects were impressive in crosses. Conclusion Wheat production is mainly affected by several environmental factors and water stress is the major constraint in Pakistan. Therefore, understanding the mechanism of plant responses to water stress is an important and basic part of crops. Thus, it is need to select such genotypes of wheat which lead to better adaptation and higher yield against water stress. These studies revealed that parental genotypes like WN-10, WN-25, WN-32 and 8126 are beneficial to generate desirable combinations and may be used in future breeding strategies. Specific combinations like, WN-35 x 8126 and WN-10 x 9526 needs further investigations. Taken together, the information derived from these studies may be used to develop drought tolerant wheat material that could give economic yield in water stressed conditions of wheat growing areas. References Ahmad M, Akram Z, Munir M, Rauf M. 2006. Physio-morphic response of wheat genotypes under rainfed conditions. Pakistan Journal of Botany 38, 1697-1702. Ahmad N, Chowdhry M, Khaliq I, Maekawa M. 2007. The inheritance of yield and yield components of five wheat hybrid populations under drought condition. Indonesian Journal of Agricultural Science 8, 53-59. Ahmadi J, Zali A, Samadi B, Talaie A, Channadha M, Saeidi A. 2003. A study of combining ability and gene effect in bread wheat under drought stress conditions by diallel method. Iranian Journal Agricultural Sciences 34, 1-8. Akbar M, Anwar J, Hussain M, Qureshi MH, Khan S. 2009. Line× tester analysis in bread wheat (Triticum aestivum L.). Pakistan Journal of Agricultural Sciences 47, 411. Akhtar L, Hussain M, Tariq A, Nasim M. 2010. A review of hundred years of wheat research and development in Punjab (1911-2010). Pakistan Journal of Science 62, 128-134. Arshad M, Chowdhry MA. 2002. Impact of environment on the combining ability of bread wheat genotypes. Pakistan Journal Biological Sciences 5, 1316-1320. Awan SI, Malik MFA, Siddique M. 2005. Combining ability analysis in intervarietal crosses for component traits in hexaploid wheat. Journal of Agriculture & Social Sciences 1, 316-317. Chowdhary M, Sajad M, Ashraf M. 2007. Analysis on combining ability of metric traits in bread wheat, Triticum aestivum. Pakistan Journal of Agricultural Sciences 45, 11-17. Day A, Intalap S. 1970. Some effects of soil moisture stress on the growth of wheat (Triticum aestivum L. Em Thell.). Agronomy Journal 62, 27-29. Hasnain Z, Abbas G, Saeed A, Shakeel A, Muhammad A, Rahim MA. 2006. Combining ability for plant height and yield related traits in wheat (Triticum aestivum L.). Journal of Agricultural Research 44, 167-1175. Hassan G, Mohammad F, Afridi SS, Khalil IH. 2007. Combining Ability in the F~ 1 Generations of Diallel Cross for Yield and Yield Components in Wheat. Sarhad Journal of Agriculture 23, 937. Hogg AC, Sripo T, Beecher B, Martin JM, Giroux MJ. 2004. Wheat puroindolines interact to form friabilin and control wheat grain hardness. TAG Theoretical and applied genetics Theoretische und angewandte Genetik 108, 1089-1097.
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