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Identification of superior parents and hybrids from diallel crosses of bread wheat (Triticum aestivum L.).

Introduction

Contributions of both i.e. nature and selective process of man have made tremendous genetic improvements in wheat plant. Increasing crop yield has still remained main concern to wheat breeders. The knowledge regarding the genetic basis of yield and its components helped wheat breeders to sort out promising parents to be used in hybridization and selection programmes. A diallel technique is one of the most powerful quantitative genetic analysis by which plant breeders estimate combining ability and heterotic potential of fixed lines or varieties. Among various diallel forms, the halfdiallel provides maximum information about genetic architecture of a trait, potentiality of parents in crosses and type of gene action controlling traits (Griffing, 1956).

The combining ability was further partitioned into general combining ability (GCA) and specific combining ability (SCA) effects; hence components of gene action were obtained. The average performance of a genotype in a series of hybrid combinations is called GCA and is used to measure additive variances and additive genes. Whereas, the performance of genotypes in crosses is determined by SCA and measures non-additive gene action (Sprague and Tatum, 1942). Importance of general combining ability for grain yield per plant was observed by Akram et al. (2011) and Khan et al. (2007) and suggested additive type of gene action for this trait. While, Shabbir et al. (2011) and Akbar et al. (2009) believed that non-additive genetic effects were high for grain yield, revealing the prevalence of SCA effects. Additive type gene action with high values of general combining ability for tillers per plant (Mahpara et al., 2008; Rahim et al., 2006), spike length (Yucel et al., 2009), spike density (Mahpara et al., 2008), grains per spike (Shabbir et al., 2011; Yucel et al., 2009) and 1000grains weight (Dhadhal et al., 2008 and Mahpara et al., 2008) were recorded. On the contrary, substantial specific combining ability effects were recorded for spike length, grains/spike, 1000 grain weight (Akram et al., 2011; Shabbir et al.,2011; Cifci and Yagdi, 2010) and spike density (Iqbal and Khan, 2006).

Proper choice of parents for hybrid wheat development is an important factor to enhance grain yield potential in existing germplasm. Nagarajan (2001) suggested the use of wheat hybrid varieties at commercial scale because of its higher production. Utilization of heterotic effects for more yield were largely attributed to cross-pollinated crops, yet evidences are now available to prove the presence of such effects in self-pollinated crops like wheat. Several plant breeders worked on heterosis in wheat and reported that grain yield can be maximized from 6% (Borghi et al., 1986) to as high as 41% (Zehr et al., 1997). Maximum and positive heterosis was found in spike length (Jaiswal et al., 2010); grains per spike (Khattab et al., 2010); 1000-grain weight and grain yield per plant (Abdel-Moneam, 2009). The objective of present investigation was to explore general combiners and mark crosses with better specific combining ability for wheat yield and its related traits by employing diallel genetic analysis.

Materials and Methods

Five bread wheat cultivars of diverse origin viz. TD-1, SKD-1, Marvi, Moomal and Mehran were crossed in half-diallel design. The seeds of parent and their 10 [F.sub.1] hybrids were sown in randomized complete block design with four replications in the experimental field, Department of Plant Breeding and Genetics. The sowing was done during mid December, 2010 with drill method and the distance between plant to plant and row to row was kept at 6 and 9 cm, respectively. The inorganic fertilizer like one bag (50 kg) of DAP per hectare was applied at the time of sowing whereas two bags of urea, one with third irrigation and second dose was applied at the time of grain formation. The crop was harvested in April, 2010. To reduce the intensity of harmful weeds, weedicide like Buctril Super at the rate of 1000cc was applied after first irrigation. The analysis of variance was carried out according to statistical methods developed by Gomez and Gomez (1984). Diallel analysis was carried out according to Griffing's Method-11, Model-1 (Griffing, 1956) by numerical approach as adopted by Singh and Choudhary (1979). Heterotic effects in [F.sub.1] hybrids were determined as the percent increase (+) or decrease (-) of [F.sub.1] hybrids over mid or better parents according to Fehr (1987) as under:

Mid parent heterosis % = [F.sub.1] - MP / MP x 100

Better parent heterosis % = [F.sub.1] - BP / BP x 100

Where, [F.sub.1] = hybrid performance, M.P. = mid parent value (the mean of both parents) B.P. = Better parent value (the mean of better parent). Ten plants per replication from each [F.sub.1] hybrid and parent were randomly selected and treated as index plants to record the data. The traits studied were tillers/plant, spike length (cm), spike density, grains/ spike, grain yield/plant and seed index (1000-grain weight in g).

Results and Discussion

The present experiment was conducted so as to estimate the combining ability and heterotic effects in [F.sub.1] hybrids developed from five parent half diallel of bread wheat. The characters studied were tillers/plant, spike length, spike density, grains/spike, grain yield/plant and seed index. The mean squares due to genotypes, parents, hybrids and parents vs. hybrids were significant for all the characters indicating the presence of considerable amount of variability in the breeding material (Table 1). These results further suggested that the parental lines and their hybrids were quite variable in their performance and presence of overall heterosis for most of the characters. The significant mean squares due to general combining ability (GCA) and specific combining ability (SCA) suggested that additive as well as non-additive genes were advocating the traits under study (Baloch et al., 2008). The magnitude of GCA variances was higher than the SCA variances indicating preponderance of additive gene effects for spike length, spike density and seed index. While high SCA variances over GCA for tillers/plant, grains/spike and grain yield/plant confirmed the prevalence of non-additive genetic effects for these traits. Importance of general combining ability variance for grain yield per plant was also observed by Akram et al. (2011) and Baloch et al.(2011) who suggested that additive type of gene action was advocating grain yield per plant, while Shabbir et al. (2011) believed that non-additive genetic effects were high for grain yield, revealing the prevalence of SCA effects. For other traits, additive type gene action with high values of general combining ability for tillers per plant was reported by Mahpara et al. (2008); spike length by Yucel et al. (2009); spike density by Mahpara et al. (2008); grains per spike by Shabbir et al. (2011) and 1000-grains weight by Dhadhal et al. (2008) and Mahpara et al. (2008). On the contrary, substantial specific combining ability effects rather than general combining ability were recorded for spike length and seed index (Akram et al., 2011; Shabbir et al., 2011; Cifci and Yagdi, 2010); spike density (Iqbal and Khan, 2006) and grains per spike (Baloch et al., 2011).

With respect to average performance, the parents TD-1 developed maximum tillers/plant, gave maximum grain yield/plant and recorded maximum seed index. While Mehran set maximum grains/spike and Marvi measured longer spikes and also recorded maximum spike density (Table 2). It is quite interesting to note that parents like TD-1 and Marvi which performed well as per se also performed well for GCA effects. These results therefore suggested that both the parents could reliably be used in hybridization and selection programmes to improve various yield traits. Regarding [F.sub.1] hybrids' performance, cross TD-1 x SKD-1 set highest grains per spike and at the same time produced maximum grain yield/plant. Hybrid TD-1 x Mehran developed maximum tillers per plant and Marvi x Moomal measured longest spikes; Marvi x Mehran recorded maximum spike density; Moomal x Mehran gave higher seed index. It is generally assumed that [F.sub.1] hybrid performance is reflected in specific combining ability, but such assumption did not hold true (Baloch and Bhutto, 2003). These results suggested that [F.sub.1] performance may not be taken as granted for specific combining ability of such hybrids. Concerning GCA effects, TD-1 expressed maximum positive GCA effects for tillers/plant and grain yield/plant. Cultivar Marvi manifested maximum positive GCA effects for spike length and spike density; Mehran for grains/spike (1.694); Moomal for seed index. These results therefore suggested that parents TD-1, Marvi, Mehran and Moomal may be preferred for hybridization and selection programmes so as to improve majority of the characters studied. The results for specific combining ability (SCA) effects revealed that maximum positive SCA effect was displayed by hybrid SKD-1 x Mehran for tillers/plant and grain yield/plant; TD-1 x Moomal for spike length and spike density; TD-1 x SKD-1 for grains/spike and Moomal x Mehran for seed index (Table 2). These results suggested that various hybrids may be considered for hybrid crop development to improve above traits.

The GCA effects presented in Table 3 revealed that for tillers/plant, TD-1 demonstrated highest positive GCA effects of 1.021 while, Mehran ranked next (0.414), yet Marvi manifested maximum negative GCA effects of -0.800. As far as SCA effect is concerned, hybrids, SKD-1 x Mehran (2.09), SKD-1 x Moomal (2.05) and Marvi x Moomal (2.01), respectively were among the top three rankers for tillers per plant (Table 4). With respect to spike length, Marvi and Moomal established maximum positive GCA effects of 1.299 and 0.406, respectively while SKD-1 exhibited highly negative GCA effect of -0.894. Eight hybrids manifested positive SCA effects while other two crosses gave negative SCA effects for spike length (Table 4). Nonetheless, top three rankers in SCA effects were; TD-1 x Moomal (1.58), Marvi x Moomal (1.19) and TD-1 x Mehran (0.83).

Regarding GCA effects of spike density, Marvi and Moomal expressed maximum positive effects of 7.158 and 2.714, respectively. Whereas SKD-1 (-6.751) and Mehran (-1.795) demonstrated negative GCA effects (Table 3). Concerning SCA effects, six hybrids manifested positive SCA effects and four crosses gave negative SCA effects (Table 4). Nonetheless, three top rankers in SCA effects were TD-1 x Moomal (11.25), Marvi x Mehran (8.92) and SKD-1 x Marvi (7.10). GCA effects of grains/spike (Table 3) revealed that Mehran and Marvi genotypes expressed positive GCA effects of 1.694 and 0.016, respectively, while TD-1 (-1.399), SKD-1 (-0.220)andMoomal(-0.091)displayed negative GCA effects. With respect to SCA effects, the top three scoring hybrids were TD-1 x SKD-1 (9.62), TD-1 x Marvi (6.28) and TD-1 x Mehran (5.61). For grain yield/plant, TD-1 and Mehran expressed maximum positive GCA effects of 1.193 and 0.371, respectively whereas Moomal (-0.736), Marvi (-0.557) and SKD-1 (-0.271) demonstrated negative GCA effects (Table 3). Concerning the SCA effects for grain yield/plant, eight crosses manifested positive SCA effects while remaining two hybrids showed negative SCA effects (Table 4). Seed index (1000-grain weight in g) results revealed that Moomal and TD-1 expressed maximum positive GCA effects of 1.215 and 1.072, respectively whereas other parents such as Marvi (-1.535), SKD-1 (-0.464) and Mehran (-0.287) displayed negative GCA effects (Table 3). From 10 crosses, eight hybrids manifested positive SCA effects yet, top three hybrids were Moomal x Mehran (4.13), Marvi x Moomal (4.12) and SKD-1 x Marvi(3.04) (Table 4).

Heterotic effects revealed that hybrids SKD-1 x Moomal exhibited maximum positive heterobeltiosis for tillers per plant; TD-1 x Moomal for spike length; Marvi x Mehran for spike density; TD-1 x SKD-1 for grains/ spike; SKD-1 x Mehran for grain yield/plant; Marvi x Moomal for seed index (Table 5). Regarding heterobeltiosis, maximum positive heterobeltiosis was displayed by hybrids SKD-1 x Marvi for tillers per plant; TD-1 x Moomal for spike length and spike density; TD-1 x SKD-1 for grains/spike; SKD-1 x Mehran for grain yield/plant and Moomal x Mehran for seed index. These crosses revealed good scope for their commercial exploitation of heterosis as well as isolation of promising progenies in later segregating generations. Analogous to our heterotic effects, Jaiswal et al. (2010) obtained mid parent and better parent heterosis for tillers per plant; Akbar et al. (2009) for spike length; Chowdhry et al. (2005) for spike density, Moneam (2009) for grain yield per plant and Akbar et al. (2009) for seed index.

Conclusion

The overall findings revealed the importance of both additive and non-additive genetic variances and effects which suggested that the use of integrated breeding strategies can efficiently utilize both gene actions. Among the parents, TD-1, Mehran, Moomal and Marvi were found as best general combiners hence can be regarded potential parents for hybridization and selection programmes to improve yield traits under study. Furthermore, if hybrid wheat becomes feasible, the better parent heterosis could be of practical value and hybrids like SKD-1 x Mehran, Marvi x Mehran, Marvi x Moomal and TD-1 x SKD-1 may be more meaningful for improving yield in wheat.

References

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Baloch, M.J., Kumbher, M.B., Jatoi, W.A., Fatima N. 2008. Estimates of genetic parameters from line x tester mating design for some quantitative traits in upland cotton, Gossypium hirsutum L. Pakistan Journal of Scientific and Industrial Research, 51: 36-41.

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Muhammad Jurial Baloch *, Toufique Ahmed Rajper, Wajid Ali Jatoi and Nasreen Fatima Veesar

Department of Plant Breeding and Genetics, Sindh Agriculture University, Tandojam Sindh, Pakistan

(received August 9, 2012; revised February 9, 2013; accepted March 6, 2013)

* Author for correspondence; E-mail:j.rrnd58@gmaiLcom
Table 1. Mean squares from diallel analysis for various yield traits
in bread wheat {Triticum aestivum L.)

Source of variation   D.F.              Mean squares
                             Tillers/plant       Spike length

Genotype              14     20.05 **            9.052 **
Parents               4      11.20 **            10.92 **
Hybrids               9      4.82 **             8.74 **
Parents vs. hybrids   1      192.53 **           4.38 **
GCA                   4      14.40 **            22.89 **
SCA                   10     22.31 **            3.52 **
Error                 42     0.477               0.524

Source of variation   D.F.               Mean squares
                             Spike density       Grains/spike

Genotype              14     334.39 **           129.69 **
Parents               4      185.44 **           113.30 **
Hybrids               9      430.21 **           28.36 **
Parents vs. hybrids   1      67.72 **            1106.56 **
GCA                   4      764.14 **           34.18 **
SCA                   10     162.49 **           167.83 **
Error                 42     34.901              0.780

Source of variation   D.F.               Mean squares
                             Grain yield/plant   Seed index

Genotype              14     18.03 **            33.57 **
Parents               4      12.12 **            52.05 **
Hybrids               9      10.62 **            16.40 **
Parents vs. hybrids   1      108.3 **            114.15 **
GCA                   4      17.40 **            36.95 **
SCA                   10     18.28 **            32.21 **
Error                 42     0.676               0.763

** = significant at 1% probability level

Table 2. Mean performance of parents and [F.sub.1] hybrids for various
yield traits in bread wheat (Triticum aestivum L.)

Parents                        Mean performance

                    Tillers/   Spike    Spike     Grains/
                    plant      length   density   spike
                    (cm)       (%)                (g)

TD-1                9.50       11.38    75.85     52.75
SKD-1               6.00       12.50    70.21     57.25
Marvi               5.50       15.75    87.05     60.75
Moomal              7.50       14.00    81.42     64.75
Mehran              8.50       13.00    72.71     65.50

[F.sub.1] Hybrids

  TD-1 x SKD-1      12.50      11.90    61.51     73.60
  TD-1 x Marvi      11.50      13.50    85.79     70.50
  TD-1 x Moomal     11.00      14.75    90.99     66.75
  TD-1 x Mehran     12.75      13.50    75.69     71.50
  SKD-1 x Marvi     9.50       14.50    85.80     68.00
  SKD-1 x Moomal    11.25      12.00    72.36     69.00
  SKD-1 x Mehran    12.00      12.75    67.41     71.00
  Marvi x Moomal    10.75      16.50    86.63     66.75
  Marvi x Mehran    11.25      15.50    92.55     71.00
  Moomal x Mehran   9.50       14.00    81.12     65.00

L.S.D (5%)          0.986      0.974    8.42      1.26

Parents                           Mean performance

                    Grain         Seed index
                    yield/plant   (g)

TD-1                19.25         45.50
SKD-1               15.50         38.75
Marvi               14.75         35.50
Moomal              16.00         40.00
Mehran              17.00         39.87

[F.sub.1] Hybrids

  TD-1 x SKD-1      21.50         42.00
  TD-1 x Marvi      20.75         41.25
  TD-1 x Moomal     19.00         44.00
  TD-1 x Mehran     19.00         40.50
  SKD-1 x Marvi     17.00         42.50
  SKD-1 x Moomal    17.50         44.00
  SKD-1 x Mehran    21.50         42.00
  Marvi x Moomal    18.75         45.25
  Marvi x Mehran    20.50         40.50
  Moomal x Mehran   18.00         46.50

L.S.D (5%)          1.17          1.247

Table 3. The GCA effects of various yield traits in bread
wheat (Triticum aestivum L.)

Parents     Tillers/   Spike    Spike     Grains/   Grain    Seed
            plant      length   density   spike     yield/   index
                                                    plant

TD-1        1.021      -0.787   -1.326    -1.399    1.193    1.072
SKD-1       -0.336     -0.894   -6.751    -0.220    -0.271   -0.464
Marvi       -0.800     1.299    7.158     0.016     -0.557   -1.535
Moomal      -0.300     0.406    2.714     -0.091    -0.736   1.215
Mehran      0.414      -0.023   -1.795    1.694     0.371    -0.287
S.E.(gi.)   0.014      0.015    0.99      0.02      0.02     0.02

Table 4. The SCA effects of.[F.sub.1] hybrids for various yield traits
in bread wheat {Triticum aestivum L.).

[F.sub.1] hybrids   Tillers/   Spike    Spike     Grains/   Grain yield/
                    plant      length   density   spike     plant

TD-1 x SKD-1        1.99       0.03     -8.79     9.62      2.37
TD-1 x Marvi        1.45       -0.55    1.66      6.28      1.91
TD-1 x Moomal       0.45       1.58     11.25     2.64      0.33
TD-1 x Mehran       1.49       0.83     0.46      5.61      -0.77
SKD-1 x Marvi       0.80       0.48     7.10      2.61      -0.39
SKD-1 x Moomal      2.05       -1.13    -1.95     3.72      0.29
SKD-1 x Mehran      2.09       0.11     -2.39     3.94      3.19
Marvi x Moomal      2.01       1.19     -1.52     1.23      1.82
Marvi x Mehran      1.80       0.69     8.92      3.71      2.47
Moomal x Mehran     -0.44      0.07     1.88      -2.18     0.15
S.E. (si.)          0.06       0.06     4.15      0.09      0.08

[F.sub.1] hybrids   Seed index

TD-1 x SKD-1        -0.05
TD-1 x Marvi        0.27
TD-1 x Moomal       0.28
TD-1 x Mehran       -1.72
SKD-1 x Marvi       3.04
SKD-1 x Moomal      1.81
SKD-1 x Mehran      1.30
Marvi x Moomal      4.12
Marvi x Mehran      0.87
Moomal x Mehran     4.13
S.E. (si.)          0.09

Table 5. Heterotic effects of [F.sub.1] hybrids over their mid and
better parents for various traits in bread wheat (Triticum
aestivum L.)

[F.sub.1]         Tillers per     Spike length     Spike density
hybrids           plant

                  R.H.    B.P.    R.H     B.P.     R.H.     B.P.
                  (%)     (%)     (%)     (%)      (%)      (%)

TD-1 x SKD-1      61.29   31.58   -0.31   -4.80    -15.77   -18.90
TD-1 x Marvi      53.33   21.05   -0.46   -14.29   5.33     -1.44
TD-1 x Moomal     29.41   15.79   16.26   5.36     15.72    11.76
TD-1 x Mehran     41.67   34.21   10.77   3.85     1.92     -0.19
SKD-1 x Marvi     65.22   58.33   2.65    -7.94    9.13     -1.43
SKD-1 x Moomal    66.67   50.00   -9.43   -14.29   -4.55    -11.13
SKD-1 x Mehran    65.52   41.18   0.00    -1.92    -5.66    -7.28
Marvi x Moomal    65.38   43.33   10.92   4.76     2.84     -0.48
Marvi x Mehran    60.71   32.35   7.83    -1.59    15.87    6.32
Moomal x Mehran   18.75   11.76   3.70    0.00     5.27     -0.37

[F.sub.1]         Grains per spike  Grain yield per    Seed index
hybrids                             plant

                  R.H.    B.P.      R.H.    B.P.       R.H.     B.P.
                  (%)     (%)       (%)     (%)        (%)       (%)

TD-1 x SKD-1      33.82   28.56     23.74   11.69      -0.30   -7.69
TD-1 x Marvi      24.23   16.05     22.06   7.79       1.85    -9.34
TD-1 x Moomal     13.62   3.09      7.80    -1.30      2.92    -3.30
TD-1 x Mehran     20.93   9.16      4.83    -1.30      -5.12   -10.99
SKD-1 x Marvi     15.25   11.93     12.40   9.68       14.48    9.68
SKD-1 x Moomal    13.11   6.56      11.11   9.38       11.75   10.00
SKD-1 x Mehran    15.68   8.40      32.31   26.47      6.84     5.34
Marvi x Moomal    6.37    3.09      21.95   17.19      19.87   13.13
Marvi x Mehran    12.48   8.40      29.13   20.59      7.47     1.58
Moomal x Mehran   -0.19   -0.76     9.09    5.88       16.44   16.25

R.H. = relative heterosis; B.P. = better parent heterosis.
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Author:Baloch, Muhammad Jurial; Rajper, Toufique Ahmed; Jatoi, Wajid Ali; Veesar, Nasreen Fatima
Publication:Pakistan Journal of Scientific and Industrial Research Series B: Biological Sciences
Article Type:Report
Geographic Code:7IRAN
Date:Jul 1, 2013
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