Analysis of variance, PCV and GCV
The pooled analysis of variance for three years of Augmented Randomized Complete Block Design (Table 1) revealed significant differences in the set of genotypes under study for all the traits except for branches per plant and days to maturity. The checks vs Genotypes component of source of variation had significant differences in terms of days to 50% flowering, clusters per plant, pods per cluster, pod length and seeds per pod. None of the traits are significant at block level.
The MSS (Table 1) of different traits at genotypic level is further confirmed by the respective PCV and GCV values. Highest estimate of GCV magnitude (reliable portion of variation) among the traits which had significant MSS over years, was recorded for pods per plant (43.73%) followed by clusters per plant (36.13%), seeds per pod (17.69%), plant height (16.66%), test weight (16.28%), pods per cluster (15.33%), grain yield per plant (14.97%), branches per plant (14.63%) and pod length (10.11%). Similar PCV and GCV magnitudes were earlier indicated for various traits in mungbean by different scientists
[Muralidhara et al., (2015), Raturi et al., (2015), Anand et al., (2016), Garg et al., (2017), Ghimire et al., (2017), Kate et al., (2017), Shiv et al., (2017) Azam et al., (2018)]. In general, the values of PCV were higher than GCV indicating the role of environment on the visible variation. These results of variance, PCV and GCV indicate ample amount of variation in the material under investigation for different traits, giving scope for utilization of the material in crop improvement programmes.
Heritability and Expected Genetic advance
The information on magnitude of GCV alone may not be trusted and these GCV values are to be viewed along with the heritability of that trait in question. Burton way back in 1952 itself emphasized the consideration of genetic variability along with heritability estimates to get an idea on expected genetic gain under selection. Highest magnitude of heritability over years was recorded in case of pods per plant followed by test weight, clusters per plant, plant height, pod length and seeds per pod. While least magnitude of heritability was recorded in case of days to maturity followed by days to 50% flowering. Low heritability was observed in case of days to maturity. While, days to 50% flowering and branches per plant registered moderate magnitude of heritability. Higher magnitudes of GCV coupled with high heritability were observed in case of pods per plant (43.73% GCV, 93.50 h
2bs) and clusters per plant (36.13% GCV, 87.76% h
2bs). The existence of higher amount of genotypic variation with high heritability will be fruitful in selection. Similarly, moderate GCV (10% to 20%) and high heritability (>60%) were recorded in case of test weight, plant height, pod length, seeds per pod, grain yield per plant and pods per cluster (Table 2).
These traits with moderate GCV and high heritability may also result in considerable gain during selection. However, it is not necessary that a trait showing higher heritability should always exhibit higher genetic advance
(Johnson et al., 1955). Therefore, estimation of expected genetic advance is important before actually planting the material for taking up selection. High expected genetic advance as per cent of mean (> 20%) was found in seven out of eleven traits, moderate (10% to 20%) in one trait (pods per plant) and low (< 10%) in two traits (days to maturity and days to 50% flowering). The characters with highest heritability [pods per plant (93.50%), test weight (89.19%), clusters per plant (87.76%) and plant height (87.56%),] have recorded highest genetic advances as per cent of means [pods per plant (87.11%), test weight (31.67%), clusters per plant (69.72%) and plant height (32.12%)]. However, the traits varied in ranking with respect to heritability and genetic advance as per cent of mean. Further, pod length which had high heritability (78.28%) recorded only moderate (18.42%) genetic advance as per cent of mean (Table 2) indicating that one can not completely rely on broad sense heritability estimates. Further, it can be inferred that considering genetic parameters separately will be always misleading hence all the genetic parameters need to be studies simultaneously for proper understanding of the trait. Higher estimates of PCV, GCV, h2bs and genetic advance as per cent of mean were recorded for pods per plant and clusters per plant. Similar higher estimates of all the genetic parameters for these two traits were also observed by
Ahamed et al., (2015) and
Muralidhara et al., (2015).
The estimates of heritability and genetic advance as per cent of mean were high for the characters
viz., plant height, clusters per plant, pods per cluster, pods per plant, seeds per pod, test weight and grain yield per plant indicating the probable operation of additive gene action in inheritance of these traits. Similar results for the above seven traits were earlier reported by
Ahmad et al., (2015); Muralidhara et al., (2015); Anand et al., (2016); Garg et al., (2017); Shiv et al., (2017); Mehandi et al., (2018); Ramakrishnan et al., (2018) and
Sharma et al., (2018). High heritability coupled with moderate genetic advance as per cent of mean was observed for pod length. Moderate heritability coupled with low genetic advance as per cent of mean was observed for days to 50% flowering. While days to maturity had low heritability and low genetic advance as per cent of mean.
Trait associations
The character associations at both phenotypic and genotypic levels over three years of evaluation with respect to eleven characters for the sixty mungbean genotypes were presented in the Table 3. From the phenotypic and genotypic correlations, it was observed that genotypic correlations are higher than phenotypic correlations. This indicates that apparent associations are largely due to genetic reasons. The traits
viz., pods per plant (0.7872**, 0.8521**), clusters per plant (0.6946**, 0.7628**), pods per cluster (0.6452**, 0.7631**), branches per plant (0.2713**, 0.3800**), days to maturity (-0.1780*, -0.2293**) and pod length (0.1781*, 0.2060**) were found to possess significant positive association with grain yield per plant in desirable direction at both phenotypic and genotypic levels. This infers that mungbean lines with higher pod number, more clusters per plant, more pods per cluster, more branches per plant and lengthier pods with less duration to maturity are productive in terms of grain yield. Further, plant height (-0.2559*) and seeds per pod (0.1494*) had negative and positive significant associations respectively, at genotypic level with grain yield per plant. Hence, selection for these two characters in the specified direction will also help in selecting genotypes with high grain yield per plant. Positive correlations are due to control of the traits which are under control of genes responsible for direct production of precursors. While, negative correlations occur due to the restricted supply of precursor for which traits compete against each other (
Madhur and Jinks, 1994). The pictorial representation of phenotypic and genotypic correlations are also indicated in Fig 1 and 2, respectively. Darker the shade stronger is the association between the traits (Fig 1 and 2).
The dependent variable also had non significant associations with days to 50% flowering (-0.0344, -0.0697) and test weight (0.0010, 0.0055), at both phenotypic and genotypic levels. This demonstrates that early flowering need not strongly associate with higher yields in mungbean despite having significant positive association between days to 50% flowering and days to maturity at both levels. The traits which are not having any significant association with yield might have assorted independently with respect to yield controlling genes. However, these traits may contribute through indirect effects of other traits.
Many researchers established associations of various quantitative characters with grain yield per plant indicating the scope of indirect selection for yield improvement in mungbean.
Ahamed et al., (2015) and
Anand et al., (2016) reported significant positive associations of pods per plant and clusters per plant with grain yield per plant at both phenotypic and genotypic levels.
Thippani et al., (2013) confirmed the significant positive association of pods per cluster with grain yield at both levels, indicating the importance of these traits during selection. The significant association of branches per plant and days to maturity with grain yield per plant in the desirable direction was reinforced by the observations of
Eswaran and Senthilkumar (2015) at genotypic level for branches per plant and
Kumar et al., (2018) for days to maturity at phenotypic level. While,
Das and Barua (2015) found significant positive association of pod length with grain yield per plant at both phenotypic and genotypic levels which is in accordance with current results.
Among various traits (independent variables), maximum association at both phenotypic and genotypic level was observed between; clusters per plant and pods per plant; pod length and seeds per pod; pods per cluster and pods per plant; days to 50% flowering and days to maturity and; pod length and test weight. Similar associations were earlier observed in mungbean by;
Anand et al., (2016) between clusters per plant and pods per plant;
Marak (2017) between pod length and seeds per pod, between pod length and test weight;
Makeen et al., (2007) between pods per cluster and pods per plant and;
Ahmad et al., (2015) and
Ramachandra and Lavanya (2017) between days to 50% flowering and days to maturity. These pairs of associations indicate that selection for one character of a pair will enable improvement in another character in that pair. This information would be helpful in improvement through indirect selection.
Direct and indirect effects of correlations
The traits, pods per plant (0.4043 and 0.3236), pods per cluster (0.3359 and 0.4579), clusters per plant (0.2071 and 0.1972), pod length (0.1130 and 0.1007) and branches per plant (0.0268 and 0.0804) had positive direct effects at phenotypic and genotypic levels and also established significant positive association at both levels with grain yield per plant (Table 4). This indicate true relationship between the above traits and grain yield per plant. Similar true relations with grain yield per plant were observed in mungbean by;
Kumar et al., (2018) for pods per plant at both phenotypic and genotypic level;
Sinha et al., (2018) for pods per cluster at genotypic level;
Anand et al., (2016) for clusters per plant at phenotypic level;
Kumar et al., (2018) for pod length at both levels;
Garg et al., (2017) at phenotypic level and
Marak (2017) at genotypic level for branches per plant. Further, days to maturity was also observed to have direct effects at phenotypic (-0.0528) and genotypic (-0.2307) levels as well as significant correlation with grain yield in negative direction. Since, early genotypes are preferred, days to maturity is to be considered to have true relation with the dependent variable and should be taken care of during selection. The true relation of these six traits is also evident from the respective R
2 values of these six traits
viz., pods per cluster (0.2168 and 0.3491), pods per plant (0.3183 and 0.2757), clusters per plant (0.1438 and 0.1529), days to maturity (0.0094 and 0.0526), branches per plant (0.0073 and 0.0306) and pod length (0.0201 and 0.0207) at phenotypic and genotypic levels respectively.
Apart from the direct positive effects, indirect positive effects through different traits are also important which actually adds up to direct effect in resulting total correlation. Such indirect positive effects were found for pods per plant (
via pods per cluster, clusters per plant, pod length, branches per plant, days to maturity and days to 50% flowering), pods per cluster (
via pods per plant, clusters per plant, days to maturity, branches per plant and plant height), clusters per plant (
via pods per plant, pods per cluster, days to maturity, branches per plant, pod length, days to 50% flowering and plant height), pod length (
via pods per plant, test weight, plant height, clusters per plant, branches per plant and days to maturity) and branches per plant (
via pods per plant, clusters per plant, pods per cluster, pod length, days to maturity, plant height and test weight) at both phenotypic and genotypic levels (Table 4). The direct effects of pods per plant, pods per cluster, clusters per plant, pod length and branches per plant on yield accounted for major portion in the total correlation. This indicate that direct selection for this trait will be effective for improvement of grain yield per plant. Further, other indirect positive casual factors of the above five traits may also be considered simultaneously along with pods per plant during selection for grain yield.
The validity of the conclusions drawn in present investigation conducted over three years is evident from the residual effect at genotypic level is 0.38 (Table 4), indicating that the characters included in present investigation had contributed around 62 percent to the dependent variable
i.e., grain yield per plant and only about 38 per cent is due to the other factors that were not considered in the present investigation.