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Vivekananda Parvatiya Krishi Anusandhan Sansthan (VPKAS) International Water Management Institute


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SABRAO Journal
of Breeding and Genetics
34(2) 95-106, 2002





Wheat (Tritium aestivum L. em. Thell) is the most important

winter cereal grown in India. During the last four decades, wheat
production and productivity increased almost six times in the
Indian subcontinent. Despite this, it is believed that wheat yields
have stagnated. To overcome this, winter wheat genes have been
introgressed into cultivated spring wheats. However, little
information is available on the combining ability of these two
distinct groups of wheat. The present study was undertaken to
determine the combining ability and gene action of yield and yield
attributes in winter x spring wheat crosses. Fifty F1 progenies
developed by crossing 10 winter and 5 spring wheat lines were
evaluated along with their parents, for yield and other attributes.
The mean squares due to female x male interactions were
significant for all the characters studied, except for days-to-
heading, biological yield/plant, and grain yield/plant. Additive
genetic effects were found to play a key role in the genetic control
of days-to-heading, plant height, and grain yield/plant. Golden
Valley and Agatha, among winter, and HD 2721 and UP 2338,
among spring wheat genotypes, were good general combiners for
most of the attributes that were studied. The cross Agent/Raj 3765
had significant specific combining ability effects for most of the
yield attributes. The estimates of specific combining ability
suggest that although general combining ability effects of most
winter wheats range from average to poor, their combination with
spring wheat parents possessing high gca can give improved

Key words: Triticum aestivum, combining ability, spring wheat, winter wheat

Wheat (Triticum aestivum L.em.Thell) is a major winter cereal crop grown in

India. By and large, wheat productivity has shown an increasing trend worldwide. During
1999-2000, India produced 75.2 million tonnes of wheat and was the second largest wheat
producer in the world, which is six-fold higher than the wheat produced in 1964-65
(Nagarajan, 2000). However, despite this huge harvest, it is widely believed that wheat
yields have attained a plateau, especially in the case of high productivity zones. Therefore,
various approaches are being followed to break this stagnation, one of which is the
introgression of winter (W) wheat genes into cultivated spring (S) wheat.

Crop Improvement Division, VPKAS (ICAR), Almora, 263 601, Uttaranchal, India;
Director, VPKAS (ICAR), Almora, 263 601, Uttaranchal, India;
* Corresponding author:

Winter wheats possess various desirable yield attribute(s), in addition to
considerable variability for profuse tillering, plant height, leaf size, maturity duration, spike
length, grain size, and grain number. They are not only responsive to high input conditions,
but have invariably demonstrated superior performance under low input conditions and a
higher degree of tolerance to a number of abiotic stresses (e.g. drought and heat) and
improved nutrient efficiency (e.g. N-efficiency and P-efficiency). Besides this, many
winter wheats show a stay-green tendency even at receding moisture levels and under high
temperature conditions (Nanda and Sohu, 1998). It is assumed that winter and spring
wheats contain different gene pools due to geographical and agroecological isolation. The
varieties Veery, Kauz, Attila, and Bavicara are good results of the utilization of the winter
wheat gene pool at the International Maize and Wheat Improvement Center (CIMMYT),
Mexico. Many promising materials developed at CIMMYT include winter wheats in their
pedigree. Presently, 80% of the advanced spring wheat lines at CIMMYT contain at least
some of winter wheat genes. It is estimated that an 8% increase in yield is due to the use of
winter wheat in the W x S hybridization programme (Nanda and Sohu, 1998).

Winter wheats have a specific vernalization and photoperiod requirement

depending upon their genetic constitution. Consequently, they do not flower in the plains
of India under natural conditions (Jag Shoran et al., 1995; Kant et al., 2001). This
peculiarity, along with a poor adaptation to Indian conditions and high susceptibility to
prevalent rust races, makes winter wheats unfit for commercial cultivation in the plains of
India. However, under hilly conditions in India, as for example Almora in the Uttaranchal
hills (29° 36′N, 79° 39′E and 1600 m asl), winter wheats usually flower without any
artificial vernalization treatment.

Yield, the most important economic character, is the product of multiplicative

interaction of various contributing characters. Hence, selection for yield per se may not be
very effective, unless its components are given due emphasis (Grafius, 1964). It is,
therefore, imperative to understand the nature of gene action and the combining ability in
winter and spring wheats for yield and yield attributes in developing high yielding
genotypes. The present investigation was undertaken to determine the gene action
governing the expression of yield and yield attributes, and to determine general and specific
combining abilities among diverse winter and spring wheat genotypes.


The 10 most important exotic winter wheat parents, namely: Agatha, Agent,
Auburn, Blueboy, Hobbit, Hustler, Orea, Wembley, Golden Valley, and Martonvasur-11
were used as female because of their good performance in their respective countries. In
addition, the five most important spring wheat parents in India, HD 2721, Raj 3765, WH
542, UP 2338, and PBW 343 were used as male, to make 50 cross combinations in a line
(L) x tester (T) mating design. These 15 parents and 50 F1s were evaluated under medium
fertility (N:P2O5: K2O:120:60:40:kg/ha) and irrigated conditions at VPKAS (ICAR),
Experimental Farm Hawalbagh, Almora, India (29° 36′ and 79° 40′E and 1250 m asl)
during the 1998-99 winter season.

The experiment was conducted in a randomized complete block design with three
replications. The plot size was 1.5 m long, one row plot, with 30 cm row spacing and 10

Table 1. Analysis of variance (mean sum of squares) for yield and other attributes in winter x spring wheat crosses.
Source d.f. Days to Days to Plant No. of No. of No. of Biological Grain 1000-
heading maturity height effective spikelets/ grains/ yield/plant yield/plant grain
tillers/ ear ear weight
Repl 2 40.96 19.47 247.88 9.87 4.74 45.60 215.41 2.51 14.60

Treat 64 83.54** 43.03** 264.86** 4.60* 5.05** 27.75** 126.98** 22.17** 54.95**

Error 128 7.21 2.82 15.92 1.46 1.57 28.80 63.80 6.69 3.65

*,**significant at 5% and 1% level, respectively.

Table 2. Analysis of variance (mean sum of squares) for yield and other attributes in winter x spring wheat crosses.

No. of
No of Grain
Days to Days to Plant effective No. of Biological 1000-grain
Source d.f. grains/ yield/
heading maturity height tillers/ spikelets/ ear yield/plant weight
ear plant.

Rep 2 40.16 21.33 198.91 7.62 5.82 45.40 185.42 2.81 21.05

Females 9 49.60** 39.70** 184.94 10.26* 4.16 363.17* 276.84** 30.34* 68.04*

Males 4 128.82** 70.13** 1399.75** 5.38 1.97 585.69** 308.03** 54.87** 76.37*

Female 36 12.46 9.83** 89.70** 3.97** 3.54** 127.23** 60.71 11.90 28.75**
X Male

Error 98 8.31 2.88 17.55 1.39 1.53 32.58 74.66 8.18 3.52

*, ** significant at 5% and 1% level, respectively.

cm plant spacing within rows. The crop received 60 kg/ha N, 60 kg/ha P2O5, and 40 kg/ha
K2O as a basal dose and 30 kg/ha as a top dressing after the first irrigation and at the
jointing stage. In addition to 304 mm rainfall, the crop received three supplemental
irrigations (50 mm, each irrigation). The crop was not protected against leaf rust (Puccinia
recondita), stripe rust (Puccinia striiformis), loose smut (Ustilago nuda tritici), and
powdery mildew (Erysphi tritici) as these diseases were minimal. However, weeds were
controlled fully and lodging prevented.

Individual competitive plants were selected randomly in each plot of three

replications for recording observations. However, the data on days-to-heading and days-to-
maturity were taken on a per plot basis. Plant height was recorded at physiological
maturity. The sampled plants were uprooted and dried. Data on effective tillers/plant,
number of spikelets/ear, and biological yield were recorded. Grain yield per plant and
1000-grain weight were recorded after threshing.

Means of cross combinations (means of five plants per replicate) for grain yield
and yield attributes were subjected to combining ability analysis (Kempthorne, 1957).
Statistical software package SPAR1 of Indian Agricultural Statistical Research Institute,
New Delhi was used for statistical analysis.


Winter x spring wheat crosses showed highly significant differences for all
characters (Table 1). Pooled analysis of variance indicated that, except for number of
spikelets/ear in male and female, both parents strongly influenced yield and yield attributes
(Table 2). Female and male interactions were evident for all parameters measured, except
days-to-heading, biological yield/plant, and grain yield/plant, indicating differential effects
for yield and yield attributes for the set of parents tested. For days-to-heading, plant height
and grain yield/plant, the larger variances in general combing ability (Vgca) compared to
those of specific combining ability (Vsca) indicated that these characters were controlled by
additive gene effects, whereas small Vgca (Vgca/Vsca <1) indicated that days-to-maturity,
tillers/plant, spikelets/ear, grains/ear, biological yield, and 1000-grain weight were affected
by non-additive genes (Table 3).

Agatha was the only female parent that exhibited positive and the highest gca for
tillers/plant, biological yield/plant, and grain yield/plant and positive gca for grains/ear,
whereas Wembley produced different gca effects for these characters. This indicated that
Agatha as female parent was better than Wembley for potential yield. Progenies of
Wembley used as the female parent showed less yield potential because of fewer
tillers/plant, spikelets/ear, grains/ear, biological yield/plant, and grain yield/plant (Tables 4
and 5). The positive gca effects for the characters except days-to-heading, spikelets/ear,
and grains/ear in the progenies of Golden Valley as the female parent demonstrated that it
was also a good female parent for yield and yield attributes. Besides this, Auburn and
Blueboy showed good general combining ability for days-to-heading, days-to-maturity, and
1000-grain weight; whereas, Agent was a good general combiner for tillers/plant,
spikelets/ear, and grains/ear. The large positive gca for tillers/plant, biological yield/plant,
grain yield/plant, and 1000-grain weight and the negative gcas for days-to-maturity
indicated that HD 2721 was a desirable male parent with high grain yield. Its progeny with
high grain yield can be screened from its crossed combinations. Except for the negative

WH 542 gca for 1000-grain weight, UP 2338 had positive gca for other yield attributes,
indicating that it is also a desirable male parent for yield. Whereas, the male parents, and
PBW 343, possessed opposite yield attributes, had negative gca effects for all the
characters, except WH 542 being positive for days-to-heading and days-to-maturity,
indicating no potential for high yield (Tables 4 and 5). Besides this, Raj 3765 was a good
general combiner for days-to-heading and 1000-grain weight but it was a poor general
combiner for plant height, grains/ear, biological yield, and grain yield/plant.

Sca resulting from non-additive genetic effects are important for the breeding
potential of cross combinations. When parents such as Golden Valley, Auburn, Agent, UP
2338, and Raj 3765 with large potential yield attributes were crossed, Agent/Raj 3765 and
Golden Valley/UP 2338 had greater scas for yield (Table 6). When parents with large gca
values for yield and its attributes such as HD 2721 and UP 2338 as males and Orea and
Wembley as females were crossed, the progeny retained high total combining ability, even
though gca effects for Orea and Wembley were small. When parents such as Hobbit and
PBW 343 with low gca for various yield attributes were crossed, the sca for yield and its
attributes was high and in the desirable direction.

Table 3. Estimation of genotypic variance, ratio of gca and sca, and predominant gene
effects for yield and their attributes in winter x spring wheat crosses.
Traits Genotypic Genotypic Ratio of Vgca Predominant
variance of gca variance of to Vsca gene effects
( σ2 gca) sca ( σ2 sca) (Vgca/Vsca)

Days-to-heading 3.40 1.40 2.42 Additive

Days-to-maturity 2.00 2.32 0.86 Non-Additive
Plant height 31.23 24.05 1.29 Additive
No. of effective
tillers/ ear 0.17 0.86 0.19 Non-Additive
No. of spikelets/ear 0.02 0.59 0.03 Non-Additive
No. of grains/ear 15.43 31.55 0.49 Non-Additive
yield/plant 10.30 4.66 0.45 Non-Additive
Grain yield/ plant. 1.36 1.25 1.09 Additive
1000-grain weight 1.93 8.41 0.23 Non-Additive


The variability currently available among spring and winter hexaploid wheats is
still extensive (Reeves et al., 1999). By introgressing genetic variability from winter
wheats, breeders have considerably augmented the yield potential of spring wheats. The
Veery wheats, developed from crosses of CIMMYT spring wheats and Russian winter
wheat, represented a quantum leap in spring wheat yield and wide adaptation. More
recently, the spring bread wheat Attila, developed from crosses with western European and
US winter wheats, had rapidly gained ground on the Indian subcontinent (Reeves et al.,

Table 4. General combining ability (gca) effects for yield and other attributes of winter and spring wheat parents in L x T crosses.
Parents Days to Days to Plant No. of No. of No. of Biological Grain 1000-
heading maturity height effective spikelets/ grains/ear yield/plant yield/plant grain
tillers/plant ear weight
Agatha -0.51 0.90* 6.05** 1.45** -0.47 4.95** 9.63** 2.29** -1.85**
Agent 0.23 1.70** 0.92 0.85** 0.73** 10.29** 1.17 0.63 -1.86**
Auburn -2.71** -1.90** 3.72** 0.12 -0.47 -5.58** -0.50 0.76 3.32**
Blueboy -3.37** -1.77** 0.65 -1.28** 0.00 -2.91* -4.43* -1.47* 3.28**
Hobbit 0.36 0.03 -3.28** 0.05 0.60* -1.45 -1.57 -0.19 -2.19**
Hustler 0.49 0.37 -6.28** -0.35 0.73** 2.29 -2.63 -1.09 -2.40**
Orea 2.09** 1.83** 0.52 0.05 0.07 1.62 0.90 0.01 0.79*
Wembley 2.29** 2.03** -1.81* -0.95** -0.67* -5.18** -4.03* -2.15** -0.26

Golden valley 0.56 -1.03** 1.39 0.59* -0.40 -3.25** 4.23* 1.89** 0.92*
Martonvasur-11 0.56 -2.17** -1.88* -0.55* -0.13 -0.78 -2.77 -0.65 0.24
SE (F)± 0.62 0.36 0.90 0.25 0.27 1.23 1.86 0.61 0.40
HD 2721 -0.57 -1.03** 6.12* 0.59* -0.17 0.52 3.73* 1.23** 1.50**
Raj 3765 -1.94** -0.03 7.12* -0.31 -0.17 -5.28** -3.10* -1.75** 1.76**
WH 542 3.49* 2.63** -9.58* -0.41* 0.03 -2.08** -3.57** -1.13** -0.94**
UP 2338 -0.04 -1.03** -1.75** 0.29 0.43* 6.75** 1.97 1.00* -0.95**
PBW 343 -0.94* -0.53* -1.91** -0.15 -0.13 0.09 0.97 0.66 -0.38
SE(M)± 0.41 0.24 0.60 0.17 0.18 0.82 1.24 0.41 0.27
*, **significant at 5% and 1% level, respectively.

Table 5. Performance of winter and spring wheat parents for yield and other attributes.
Parents Days to Days to Plant height No. of No. of No. of Biological Grain 1000-grain
heading (Days) maturity (cm) effective spikelets/ grains/ ear yield yield/plant weight (gm)
(Days) tillers/plant ear (gm) (gm)

Agatha 128.67 A 165.67 P 76.67 P 6.33 G 21.67 A 78.67 G 25.33 G 10.30 G 35.73 P

Agent 126.67 A 165.67 P 78.67 A 5.67 G 20.67 G 56.67 G 24.67 A 10.37 A 46.53 P

Auburn 127.67 G 165.00 G 67.00 P 6.00 A 23.00 A 68.33 P 18.00 A 6.80 A 32.07 G

Blueboy 130.67 G 166.00 G 74.67 A 4.33 P 21.33 A 61.33 P 16.67 P 7.00 P 33.90 G

Hobbit 129.67 A 164.67 A 74.67 G 5.33 A 20.33 G 47.33 A 17.33 A 8.10 A 39.43 P

Hustler 141.00 A 173.33 A 102.67 G 6.33 A 20.00 G 48.33 A 22.00 A 6.40 A 40.10 P

Orea 147.33 P 178.67 P 94.33 A 8.33 A 21.33 A 49.33 A 25.33 A 7.60 A 36.30 G

Wembley 148.33 P 78.67 P 70.33 G 5.67 P 18.67 P 43.00 P 10.00 P 4.30 P 29.53 A
Golden 143.67 A 175.33 G 85.00 A 7.00 G 21.00 A 57.67 P 20.00 G 7.03 G 32.00 G

Martonva 146.33 A 176.67 G 79.67 G 5.33 P 19.33 A 52.00 A 15.33 A 6.40 A 31.90 A

HD 2721 146.33 A 177.00 G 83.00 P 5.67 G 20.00 A 56.00 A 22.33 G 8.03 G 30.77 G
Raj 3765 146.00 G 178.23 A 91.33 P 7.00 A 21.33 A 44.33 P 22.00 P 6.43 P 33.37 G
WH 542 151.33 P 179.67 P 63.67 G 6.67 P 22.00 A 62.67 P 23.33 A 7.17 P 37.53 P
UP 2338 139.67 A 173.00 G 66.67 G 6.00 A 22.33 G 72.00 G 16.67 A 6.57 G 26.47 P
PBW 343 147.67 G 178.33 G 71.67 G 5.67 A 17.33 A 43.00 A 163.67 A 6.60 A 42.43 A

A= Average, P= Poor, G= Good

Additive gene effects were found in days-to-heading, plant height, and grain
yield/plant. As this is a fixable component, selection is expected to bring substantial
improvement in these characters. Nanda et al. (1990) suggested the importance of both
additive and dominant gene effects for days-to-heading and plant height in the F2 of a
winter x spring wheat cross. Prevalence of additive genetic effects in winter x spring wheat
crosses were also suggested by Sharma et al. (1995) for plant height, Kant et al. (2001) for
days-to-heading, plant height and spikelets/ear. On the other hand, non-additive gene
effects were predominant for days-to-maturity, tillers/plant, spikelets/ear, grains/ear,
biological yield, and 1000-grain weight from the analysis of variance components for
combining ability. Salgotra et al. (1997) and Kant et al. (2001) also reported non-additive
genetic control of tillers/plant, grains/ear, 1000-grain weight, and biological yield/plant in
winter x spring wheat crosses. Thus, it would be important to evaluate not only the gca of
parents, but also the sca of the cross combinations.

Analysis for combining ability demonstrated that gca for Golden Valley as the
female parent was greater than that of UP 2338 for the potential quantum yield (Tables 4
and 5). When Golden Valley winter wheat was used as the female parent and UP 2338
spring wheat as the male parent, their progeny had high and significant sca effects for lower
plant height and higher biological yield, grain yield, and 1000-grain weight (Table 6).
These parents were good general combiners for these traits. These results also indicated
that the parents included in this study were quite genetically diverse and their diverse genes
have combined to cause these effects. This cross is valuable because of the presence of an
additive gene action and may respond to conventional selection methods. As expected
from their parental values, the winter wheat line Agent would be the best genotype to use as
a parent for developing progenies having fewer number of days-to-heading, higher grain
yield, and 1000-grain weight (Table 5). Moreover, Agent also contributed to effective
tillers/plant, spikelets/ear, and grains/ear. Its progeny with high grain weight, yield, and
other attributes can be screened from its crossed combinations. The spring wheat variety
Raj 3765 combined with Agent to produce progeny with early days-to-heading and higher
tillers/plant, spikelets/ear, grains/ear, and 1000-grain weight (Table 6), although gca effects
were good, average and poor for these characters in one of the parents. These results
indicate that yield and yield attributes were affected by genes from both female and male
parents and additive gene action was important in the parents having high gca, which
complement with the genes of a low gca parent. In such a situation, desirable transgressive
segregants can be obtained by crossing high x low gca combiner and in selecting F2
generation (Langham, 1961). In the present study, when parents such as Hobbit and PBW
343 with average or poor gca effects for days-to-heading and tillers/plant, grains/ear, and
grain yield/plant were crossed, the F1 showed a greater sca for these traits. Orea combined
well with HD 2721 as their F1 had early days-to-heading, higher spikelets/ear, grains/ear,
and 1000-grain weight, though the gca effects of both parents for these traits were average,
except for 1000-grain weight. It may be inferred that diverse gene constellations for these
traits would have caused this effect (Langhum, 1961). Based on these results, it may be
inferred that both per se performance as well as gca are important in the selection of

Among the spring wheat parents, HD 2721 and UP 2338 provided large gca for
yield and yield attributes (Tables 4 and 5). Raj 3765 and WH 542 provided intermediate
gca values, whereas PBW 343 had the smallest gcas for yield and yield components.
Although PBW 343 is one of the most widely cultivated varieties in India with high yield

Table 6. Crosses having significant sca effects in L x T winter x spring wheat crosses.
Crosses Days to Days to No. of No. of No. of Biological Grain 1000-
heading maturity effective spikelets/ ear grains yield/plant yield/plant grain
tillers/ plant /ear weight

Agent/Raj 3765 - 0 + + + 0 0 +

Orea/HD 2721 - 0 0 + + 0 0 +

Hobbit PPW 343 - 0 + 0 + 0 + -


Golden 0 + 0 0 0 + + +

Wenbley/ UP 0 0 0 + + 0 0 0

Golden 0 - - 0 0 0 - +
Valley/HD 2721

Auburn/ WH 542 0 0 + 0 0 + + -

(-) = significant in negative direction, (+) = significant in positive direction, (0) = not significant.

levels and adaptability, no valuable cross combinations were obtained with this set of

These results suggest that breeders have been able to manipulate such yield
attributes as tillers/plant, plant height, spikelets/ear, grains/ear, grain yield/plant, and 1000-
grain weight in the pursuit of obtaining higher yielding genotypes both in spring as well as
in winter wheat. Raj 3765, a released variety, has a higher number of tillers/plant. Agent
had higher grain yield/plant and 1000-grain weight. In contrast, Wembley had the lowest
grains/ear, biological yield, and 1000-grain weight making it unfit for cultivation in Indian

Cross combinations of Agent/Raj 3765 showed significant sca effects for most of
the yield contributing characters followed by Hobbit/PBW 343, Orea/HD 2721, and Golden
Valley/UP 2338. In addition, combining ability estimates of the parents revealed the
importance of both additive as well as non-additive gene effects. Although most of the
parents’ gca were average, nevertheless, possibilities exist to develop desirable genotypes
by involving appropriate parents in a crossing programme. These results while restricted to
these specific parents suggest that multiple crossing followed by conventional selection
may improve yield attributes and, consequently, yield.


The authors acknowledge the help of Mr. B. D. Pandey and Mr. Daya Shanker
during the course of the field experimentation and data recording.


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