Direct and indirect selection for yield in chickpea

Summary The efficiency of indirect selection for seed yield was compared with direct selection for yield per se in chickpea. A total of 2500 single F₂ plants, derived from 50 crosses with 50 plants from each cross, were divided into five sub-populations (SP1 to SP5) of 500 plants each by including 10 plants from each of the 50 crosses. The five sub-populations were advanced upto F₆ by exercising 10% selection intensity for four successive generations for number of pods per plant in SP1, number of seeds per pod in SP2, seed weight in SP3, seed yield in SP4 and random selection in SP5. The efficiency of direct and indirect selection for yield was evaluated by comparing groups of 50 F₆ lines from each sub-population. SP1 and SP3 F₆ lines showed higher mean grain yield than the other three methods. SP1 and SP3 were found to be almost equally efficient in developing F₆ lines which were significantly superior to the check. This suggests that indirect selection for yield via pod number and seed weight is more efficient than direct selection for yield.     

Intra- and intergeneration relationships among yield,its components and other related characteristics in spring wheat

Summary The relationships between yield, its components and other associated characteristics, both within and across generations, were studied in the F₂, F₃ and F₄ of two Hard Red Spring wheat (Triticum aestivum) crosses using simple correlation, path coefficient and step-wise multiple regression analyses. In the F₂ and F₃ the plants were grown 50 cm apart from each other while in the F₄ they were grown under the usual farm practices. Selection was practiced for high and low yield in the F₂ and F₃ mainly on the basis of individual plant yield. Statistically significant, but not always practically useful, correlations were found between yield and its components and other associated characters. The relationship between yield and protein content was negative and significant within all generations but not so between F₂ (and F₃) and F₄. The intergeneration correlation coefficients between F₄ grain yields and grain yields measured in the F₂ and F₃ were all positive and highly significant. These coefficients, which are also heritability estimates in standard units, were small in magnitude.Stepwise multiple regression analysis identified plant yield as the most significant factor in determining F₄ line yield, followed by its components in the order of 1000-kernel weight, grain yield per plant and number of tillers per plant.Path coefficient analysis identified tiller number per plant and grain yield per spike as having strong positive direct effects on single plant yield. Harvest index of individual F₂ plants can be used as an indirect selection criterion for yield.     

Evaluation of Early Generation Testing in Chickpea1)

Inter-generation correlations between F₂-F₃, F₂-F₄ and F₃-F₄ in six crosses of chickpea were calculated by using individual plant/progeny means. In general, correlation values in case of plant height, seeds per pod and 100-seed weight were higher between F₃ and F₄ than those between F₂ and F₃ and F₂ and F₄. However, inter-generation correlations were mostly non-significant in case of pods per plant and grain yield. These results clearly show that selection in F₃ can be fruitful for seeds per pod and 100-seed weight. However, selection for pods per plant and grain yield in early generations may not show any relationship with later generation performance for these traits.     

Single and Multiple Traits Selection in a Segregating Population of Wheat, Triticum aestivum L.

The breeding materials used in this study were the F₃, F₄ and F₅-generations of the cross between Giza 158 × Sonora 64 (Triticum aestivum L.). The objective of this study was to compare the relative merits of Smith-Hazel, desired gain selection indices, independent culling levels and single trait selection in improving grain yield, heading date and several agronomic traits. Highly significant differences among F₃ families and a satisfactory genotypic coefficient of variability were obtained for all the traits studied. The genotypic correlations were high between yield and each of spike weight, kernels/spike and spikes; plant, intermediate with 1000 kernel weight and very low with heading date, plant height and spike1 length-After two cycles of selection, the results of the gains realized indicated that the most effective method for improving yield was the Smith-Hazel index (SH₇) of seven traits followed by the desired gain index of seven traits (DG₇), SH₅, independent culling levels, DG₅ and direct selection (or grain yield/plant. Direct selection for heading date, plant height and spike length was the best method for improving these traits, but undesirable correlated responses in the other traits were obtained.     

The relationships between plant form,competitive ability and grain yield in a barley cross

Summary The grain yield of F₅ lines from a barley cross, grown in field plots, showed no correlation with F₃ single plant grain yield. F₅ grain yield did however show a significant inverse relationship with plant height and leaf length in the F₃, especially at high levels of nitrogen. Shorter plants with shorter leaves in the F₃ tended to produce lines of similar habit in the F₅, and these characters were associated with lower F₃ grain yield and higher F₅ grain yield.These relationships are believed to depend on the different circumstances of competition in the F₃ and the F₅ respectively. It is concluded that plant form in segregating generations of cereals may provide a valuable selection criterion for high grain yield in monocultures.     

Efficiency of Early Selection for Yield Performance in Wheat

Selection for yield in early generations is difficult, mainly due to a shortage of seed. In the breeding company Fr. Strube the breeder uses three-way crosses and first selects for yield between F₂′-derived lines in the F₄′. The generations are called F₁, F₂′, etc., because the first generation is already segregating. The efficiency of this selection was checked in the F₅′ and F₆′. For this purpose F₃′-derived progenies, corresponding to poor and good yielding bulks in F₄′, were selected. In later generations the average yield of the group of strains with good yield in the F₄′ was 4 % (F₅′) and 3 % (F₆′) higher, respectively, than the mean of the strains with low yield in the F₄′. Strains with very low yielding potential can easily be discarded in the F₄′. The risk of discarding lines with a very high yield is minimal. Selection for yield in the F₄′ was as effective as in the F₅′. The importance of testing F₂′-derived bulks in the F₄′ is not only based on the assessment of yield but also on the possibility of reliable intensive selection through recording visual characters (height, resistance against diseases) in a second environment in parallel to the single plant progenies.     

Effect of selection for pollen grain size on various traits in common bean

Summary Selection among microgametophytes usually exploits variation in pollen grain germination. Studies of variation in pollen grain size in common bean (Phaseolus vulgaris L.) suggested that selection for size might lead to changes in sporophytic traits. To determine whether microgametophytic selection based on size would affect pollen grain size in subsequent generations or sporophytic traits that were correlated with pollen grain size, pollen grains from three crosses were separated into two size categories by sieving and then used to pollinate cv. Diacol Calima. Selection resulted in changes in pollen grain diameter for pollen from F₁, F₂ and F₃ plants for all crosses. In vitro germination indicated no differences between vigor of large and small grains, but extraction and sieving reduced germinability. F₁ seed from two of the crosses with size-selected pollen varied in weight according to pollen grain size, but in subsequent generations, the effect disappeared. Both size categories of selected pollen resulted in F₂ progeny with reduced numbers of seeds per pod as compared to controls, suggesting that the size selection process may have resulted in indirect selection for traits reducing seed set. The overall results suggested that genes determining pollen grain size in bean have little or no effect on sporophytic traits such as seed size and seed yield.     

Response to early generation selection for grain yield and harvest index in bread wheat (T, aestivum L.)

The availability of a reliable selection criterion for the identification of the most productive genotypes in early generations represents a crucial point in many breeding programs. The findings reported in the literature concerning the effectiveness of early generation testing for grain yield (GY) or harvest index (HI) are contradictory. In this work, we measured the response to selection for GY and HI applied in different generations, from F₂ to F₆ in nine segregating populations previously screened in F₂ by means of visual selection. Genetic variability for HI and GY was found in most of the segregating populations. However, GY of spaced plants in F₆ and F₃ generations was weakly correlated with yield of F₄ and successive generations grown at normal seed density. HI was of limited value as an indicator of yield potential.     

The effectiveness of early generation (F3) yield testing and the single seed descent procedures in two cowpea (Vigna unguiculata (L.) Walp.) crosses

Summary Two breeding procedures were compared in two cowpea crosses. Both procedures were started from the same selected F₂ plants in each cross. For the early generation yield testing, the F₃ lines were yield tested and a pedigree and bulk methods followed in F₄ and F₅. Each F₅ line was bulked to provide seed for a yield test in F₆. In the Single Seed Descent (SSD) procedure, a single seed was taken from each F₂ plant to produce the F₃ generation. The procedure was repeated for the F₄ and F₅. The F₆ SSD lines and the F₆ yield testing lines were compared in yield tests.The results showed that differences in yielding ability of F₃ lines persisted over generations indicating that selection was effective. This was confirmed by the high significant correlations between F₃ yields and those of later generations which ranged from r=0.51^* to 0.85^**.The grain yields of lines derived by the single seed descent procedure were as good as those derived from early generation yield testing.Significant linear correlation between visual rating of F₃ and F₆ yields with actual yields revealed that it is possible to identify promising lines of cowpea visually.     

Combining ability of soybean (Glycine max) for low phosphorus tolerance on acidic soils of Western Ethiopia

Determining the gene actions governing the inheritance of traits of interest has paramount importance in designing a breeding approach to improve the progeny populations. This study was undertaken to determine the combining ability of nine selected parental lines in the F₂/F₃ segregating populations for low P tolerance. The experiment was laid out in an alpha lattice design in two locations, that is Mettu and Assossa of Western Ethiopia that are characterized by soils with low P and pH. General combining ability (GCA) effects were highly significant for grain yield, pod length, days to maturity and plant height, while specific combining ability (SCA) effects were highly significant for grain yield, 100-seed weight, pod length and plant height. The parent Hardee-1 was identified as the best general combiner for yield, number of seeds per pod, pod length, plant height and pod number. The results suggest that additive gene action was important for several of the studied traits, implied by significant GCA effect, which might indicate selection for these traits could be effective in later segregating generations.     

Selection response for carbon isotope discrimination in a Triticum polonicum×T. durum cross: Potential interest for improvement of water efficiency in durum wheat

This study has been conducted to evaluate the usefulness of carbon isotope discrimination (δ) in mature kernels as a criterion for the improvement of water-use efficiency and yield under drought in durum wheat. For this purpose, Triticum durum‘Om Rabi 5’ was crossed with T. polonicum pseudochrysospermum 9 (Tp9) which has been found to be more drought tolerant and to have a lower carbon isotope discrimination value of the grain. The F₂ population showed a wide segregation for carbon isotope discrimination. Further, divergent selections (selection of plants most different in carbon isotope discrimination) were made among individual F₂ plants, and for carbon isotope discrimination in F₃ progenies under field conditions. Selected F₃ and F₄ progenies were evaluated under field conditions for morphological and agronomical traits. Broad-sense heritability (h²_b), response to selection and realized heritability (h²_r) were high. The narrow-sense heritability (h²_n= 0.37 ± 0.047) indicated that additive and dominance effects were involved in the genetic control of carbon isotope discrimination. Negative correlations were noted between carbon isotope discrimination and grain yield and between carbon isotope discrimination and biomass yield within years and generations. An explanation of this result is attempted by analysing the relationships between carbon isotope discrimination and several phenological and morphological traits influencing the water-use efficiency. The divergent groups selected for low and high carbon isotope discrimination exhibited significant differences for days to heading, plant height, shape of the spike and number of spikelets per spike. Correlations were also found between carbon isotope discrimination and plant height, harvest index, shape of the spike, spike length, and number of spikes per plant. The potential use of carbon isotope discrimination as a criterion for the improvement of water-use efficiency in durum wheat is discussed by considering the genetics of this trait (variability, heritability, response to selection) and also the associations with phenological and morphological traits.     

Why SR52 is such a great maize hybrid? II. Heritability,correlation and path-coefficient analyses

Although exceptional heterosis for yield in the world’s first commercial single cross maize (Zea mays L.) hybrid SR52 has been confirmed, the role of secondary traits hasn’t been established. Objectives of this study were to establish relationship between grain yield and secondary traits, while assessing heritability and genotypic variation in SR52’s segregating populations. Knowledge on associations between yield and secondary traits would be crucial in extracting productive inbred lines from the population. Traits were subjected to correlation and path-coefficient analyses. Path analysis model accounted for over 70% of variation (R²) in all populations. Consistently high positive correlations with grain yield were observed for ear length and number of kernel rows on ear. Ear girth (0.19), and number of kernels per row (0.48) showed high direct effects on yield in the F₂ population and the trend was consistent in the backcrosses. Positive indirect effects of most traits on grain yield were negligible with the exception of kernel per row via total number of kernels on ear, which was above 0.40 in the F₂ and BCP1. Ear length can be exploited for indirect selection for yield in SR52’s segregating generations. 100-kernel weight and number of kernel rows on ear can, to a lesser extent, be targeted in selecting for grain yield. High to medium phenotypic variability in segregating generations of SR52 for traits such as ear length, number kernel rows on ear and grain yield should give impetus to selection of productive inbred lines from SR52 hybrid because of presence of diversity in the gene pool.     

Comparison of pedigree selection and single seed descent for oil yield in linseed (Linum usitatissimum L.)

Summary Four populations of linseed derived from five parents were advanced from the F₂ to the F₆ generation by SSD and to the F₅ by pedigree selection. These populations were used to compare the efficiency of the two methods in order to provide superior genotypes with respect to grain yield and oil content, i.e. oil yield. The results showed minimum differences between pedigree and SSD lines for grain yield, where in only one cross the SSD lines were significantly superior to the pedigree lines. Since pedigree selection was carried out for both, seed yield and oil content, a positive response to selection was expected. However, early selection for yield, a character with low heritability, was not successful. On the contrary, significant differences for oil content were detected between the two groups of lines in three of the four crosses studied. In these cases the pedigree lines were superior to the SSD lines. These results demonstrate that strict selection in early generations for oil content, a character with comparatively high heritability, is feasible and successful in linseed. However, selection for seed yield should be postponed to later inbred generations. Consequently, in breeding for maximum oil yield of linseed a two-step selection procedure is recommended.Abbreviations PS pedigree selection - SSD single seed descent - TGW thousand grain weight     

Selection in Early Generations of Dried Peas, Pisum sativum L. II. Significance of the Environment of Selection

In early generations peas arc frequently grown widely-spaced against wire netting under growing conditions considerably different from the commercial cultivation practice of dense stands. In 1985 and 1986, five morphologically different pea varieties were grown both widely-spaced against wire and in dense stand in plots. In the plot stands no methods to prevent the peas from lodging were used. Grain yield and yield-related traits were recorded in both environments of selection, i.e. ‘wire netting’ and ‘plot stand’. Data were analyzed with respect to the suitibility of the environment ‘wire netting’ for selection for grain yield to be realized in commercial stands of the crop. Against wire, the grain yield, number of podded lateral branches, number of podded nodes/main stem, and the profile of grain yield/node on the main stem were the most responsive characters as compared to the standard plot with a dense stand. The relative increase differed between genotypes, which was also indicated by highly significant genotype × environment interactions. Only small changes consistent over genotypes were observed for thousand grain weight (slightly lower), number of seeds/pod (slightly increased) and grain yield/pod (slightly higher). Against wire, the expression of varietal variation was higher for most characters. In addition, single plant error was lower, so that at the wire heritability values based on individual plants were larger than in the plots. The significance of the present results and the effects of competition on selection in early generations in dried peas are discussed.     

Selection for Fusarium head blight resistance in early generations reduces the deoxynivalenol (DON) content in grain of winter and spring wheat

Fusarium head blight (FHB) results in yield losses and contamination of kernels by mycotoxins, particularly deoxynivalenol (DON). For minimizing DON content in grain, indirect selection methods would increase gains from selection compared to the costly and time‐consuming DON analysis. The aim of this study was to examine whether an early selection for fewer FHB symptoms would lead to a reduced DON content in grain after inoculation with Fusarium culmorum. Starting with a double‐cross derived population of about 1,100 genotypes, 30 F_1:3 genotypes were selected for FHB rating in a two‐step selection in spring wheat with the non‐adapted resistance sources CM82036 and ‘Frontana’. In winter wheat, 30 F_1:2 genotypes were selected out of a double‐cross derived population of about 600 F₁ plants from crosses with German resistance sources (‘Dream’, G16‐92). Selected genotypes were grouped in three categories according to their FHB rating (low, moderate and high) and analysed afterwards for grain DON content. The three groups differed in their DON content illustrating that indirect selection should already be feasible in the earliest generations. Because of the wide genotypic ranges for DON contents within one grouping, a final DON analysis for selected materials is advisable to achieve full selection gain.     

Prediction of F6 line performance from F3 families in spring oilseed rape. I. Yield and other agronomic traits

Summary The reliability of a selection among crosses based on a cross prediction in early generations was investigated in spring rapeseed. The performance of the parents, the F₂ generation, and random F₃ lines from four crosses were used to predict the probability of finding superior recombinant lines. These predictions were made for two years and compared with the observed performance of F₆ lines in the second of these two years and in an additional year. Predicted and observed performances coincided reasonably for the characters plant height, standability, maturity and an index calculated from seed yield, oil content and protein content. For seed yield and flowering time, the predictions were very unreliable. In conclusion, prediction methods may be useful in rapeseed breeding, if quality traits are of major commercial interest.     

Selection for biomass yield in wheat

Summary Biomass (above ground plant parts) yield may be a useful selection trait for yield improvement in wheat (Triticum aestivum L.). This study was conducted to estimate realized heritability of biomass yield and to determine the response to selection for high and low biomass yield in 8 genetically diverse populations of spring wheat under two production systems. Selections were made among the F₃ lines. Progenies of the selected lines were evaluated in replicated field tests in the F₄ generation under high fertility and low fertility production systems at Rampur, Nepal, in 1991. Fertility level had a significant effect on biomass yield, grain yield, effective tiller number, number of kernels per spike, thousand kernel weight, and harvest index. Selection in the F₃ for high and low biomass yield was effective in identifying F₄ lines with high and low biomass yield, respectively. Biomass yield differences between high and low selection groups in the F₄ generation, expressed as percent of the mean of the low selection group and averaged over the eight populations, were 53.9 and 36.5% higher than the mean of the low selection group under the high and the low fertility production systems, respectively. The corresponding figures for grain yield were 48.8 and 34.9% under the high and the low production systems, respectively. Also, selection for high biomass yield resulted in higher effective tiller number, and number of kernels per spike, but lower harvest index. Realized heritability estimates for biomass yield were greater at high fertility (range 0.49 to 0.85) than at low fertility (range 0.22 to 0.44). Biomass yield showed positive genotypic correlations with grain yield, effective tiller number, and number of kernels per spike but a negative correlation with harvest index. The results indicated that selection for high biomass yield should bring about positive improvements in biomass yield, grain yield, effective tiller number, and number of kernels per spike. The correlation between F₃ and F₄ generations suggested that biomass yield in the F₃ generation was a good predictor of biomass yield and grain yield in the F₄ generation. Selection for biomass yield in wheat should be made under the standard production system to obtain a realistic response.     

Honeycomb selection for yield in early generations of spring wheat

Summary The effectiveness of the honeycomb selection method for yield in spring wheat (Triticum aestivum L.) was evaluated using progenies from two wheat crosses, Glenlea x NB131 and Glenlea x Era. Honeycomb selection was carried out in the F₂ and F₃ generations, grown at the University of Manitoba in the summers of 1980 and 1981, respectively. In both generations, divergent selection was made for both high and low yield. Plants selected in the F₃ generation were entered in an F₄ yield test in the summer of 1982. Results of the experiment showed that honeycomb selection for yield in the F₂ and F₃ generations was effective in identifying parents of high- and low-yielding lines. F₃ plants from highyielding F₂ selections gave higher yields than those from low-yielding F₂ selections by 11.5% and 13.0% for Glenlea x NB131 and Glenlea x Era crosses, respectively. The F₄ yield test showed that high yielding selections from both crosses significantly outyielded by 8.9% low yielding selections and by 14.4% the unselected composite lines. It is concluded that the honeycomb selection method can be used for early generation selection in spring wheat.     

Selection for Seed Yield in Lolium perenne L.*

Components of seed yield were estimated for each set of 100 plants derived from six varieties of Lolium perenne. Significant differences between the varieties were found for all components of seed yield, but the range was broader within each variety. An indirect selection of seed yield using the character seeds per spike produced a selection response similar to that obtained by direct selection on high seed yield per plant. A direct and indirect selection of characters estimated in small, drilled plots of topcross off-springs (F₁) was less successful than the selection based on the single plants. Spearman rank correlation coefficients between the single plants and their offsprings did not show significant agreement for seed yield. But altogether the results indicated that the increase of seed yield/area was about 6 % when 10 % of the best plants were selected.     

Studies on the breeding of self-pollinating cereals

Summary The limitations of the pedigree and mass methods of selection in the improvement of self pollinating cereal crops are discussed and the value of quantitative data in the early generations of selection is emphasised. Two systems of selection which provide such data have been introduced at the Cambridge Plant Breeding Institute.In one, known as the F₂ progeny method of selection, the progenies of selected F₂ plants are grown in yield trials without further selection during F₄, F₅ and F₆ after which single plant selections are made within the more promising progenies.In the other system, known as the pedigree trial method of selection, normal pedigree selection is carried out in F₂ and F₃. In the F₄ generation single plants are selected from the better families for continued pedigree selection and within each of these families the remaining plants are bulked to give grain for a yield trial in the following year. This process is repeated in F₅ and F₆, the grain for trial being in each case obtained from the progeny rows and not from the preceding trial.The use of these methods is illustrated by reference to wheat hybrids grown at Cambridge, and the data obtained is shown to be useful in comparing the value of different parental combinations as well as in assessing the relative merits of the selections within each cross.