Genetic response to index selection for grain yield, kernel weight and per cent protein in four wheat crosses

Quantitative genetic theory was used to investigate selection differentials, expected and observed direct and correlated responses to simultaneous improvement of grain yield, kernel weight, and grain protein content in F₃ and F₄ populations of four spring wheat (Triticum aestivum L.) crosses. Selection in the F₃ generation based on the Smith-Hazel index (SH) and yield was found to be superior to the other methods studied in identifying high-yielding lines, but resulted in substantial decrease in grain protein level. Consequently, a 1.0% increase in protein from selection for protein depressed grain yield as much as 536 kg/ha below the population mean, reducing the expected yield gains per generation selection cycle by 250%. The weight-free indices (EW), particularly the index involving only yield and kernel weight (EW3) and its linear approximation (EW4), and the desired gains indices were effective in improving protein content but were less efficient in selecting top-yielding lines. Selection in the opposite direction using truncation of the lowest 10% of the population based on weight-free-index involving yield, kernel weight and protein (EW2) reduced all traits significantly compared with the mean of the unselected population indicating the effectiveness of the unweighted selection index. The observed genetic gains (Ra) from selection based on yield, the SH, EW3 and EW4 indices were slightly lower than the expected advances per cycle (R) in populations from crosses ‘Sinton’בGlenlea’ (C1), ‘Glenlea’בNB505’ (C2), and ‘A2P5’בNB320’ (C6) (Ra/R = 70 to 85%) but were higher in cross ‘NB505’בA2P5’ (C5) (Ra/R = 126 to 143%). It was concluded that weight-free and the desired gains indices can be used to improve wheat grain yield and grain protein simultaneously in F₃ generation selection, as revealed by response to selection measured in the F₄ generation.     

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.     

Experimental evaluation of several cycles of marker-assisted selection in maize

A program was initiated in 1994 to compare the efficiency of marker-assisted selection (MAS) and conventional phenotypic selection. A population of 300 F_3:4 families was generated from the cross between two maize inbred lines F2 and F252 and selected on an index combining grain yield and grain moisture at harvest. This population was characterised for 93 RFLP markers and evaluated as testcrosses in a large range of environments. Three methods of selection were applied (i) two cycles of conventional phenotypic selection; (ii) two cycles of MAS based on an index combining phenotypic values and QTL genetic values and (iii) one cycle of combined MAS followed by two cycles of selection based only on the QTL effects estimated in the first generation. The different populations were characterised for RFLP markers. The evolution of allele frequencies showed that selection on only-markers was very efficient for fixing QTL alleles found favourable in the initial population. This evolution was quite different from that observed for phenotypic selection or combined MAS. Genetic gain was evaluated and found significant for each method of selection. Nevertheless, the difference between phenotypic selection and combined MAS was not significant. The two additional cycles of MAS on only-markers did not improve significantly the genetic value of the population. Moreover, the genetic variance of this population remained high, despite most of the QTL initially detected were almost fixed. The results suggest that the QTL effects estimated in the initial population were not stable due to epistasis and/or QTL by environment interactions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Breeding for yield and quality in durum wheat

Three populations of an intervarietal durum wheat cross IWP5308/PDW208, F₅, F₅BIP₁ (population derived after intermating in F₂) and F₅BIP₂ (population derived after intermating in BIPF₁), were evaluated under three different agronomic environments for mean performance and stability of genotypes for grain yield, yield components and protein content. Though the biparental progenies indicated a higher mean performance, they did not differ significantly from progenies of the pedigree method for almost all characters. The biparental progenies, however, produced a higher number of stable genotypes for grain yield per plant, grains per ear and protein content. The F₅ population had a higher number of stable genotypes for 1000 grain weight and number of tillers per plant. The BIP progenies also had a higher number of genotypes with above average mean performance, and many were significantly higher than the checks WH896 and WH542, compared with F₅ progenies. Hence, in spite of high G x E interactions, the use of cycles of biparental mating and selection of top yielding lines on the basis of yield components can enable selection of stable genotypes with high protein content. Number of tillers per plant and 1000 grain weight were the yield component characters which made maximum contribution to phenotypic stability of the genotypes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Improvement of maize populations for resistance to downy mildew

Upgrading levels of disease resistance are a primary objective of maize breeding programmes. Efficacy of S₁ recurrent selection in improving levels of resistance to downy mildew (DM) infection was assessed in Nigeria from 1997 to 2000 in six maize populations. Improvement procedures consisted of evaluating S₁ progenies under artificial infection with DM spores and in disease‐free environments and using a selection index to combine selection for reduced DM infection with appropriate agronomic characters from more than one environment. Three to four cycles of selection were completed in each of the populations. Products from the different cycles of selection were evaluated and data collected on DM infection parameters and agronomic traits. Result obtained showed 3–4 cycles of selection were adequate to reduce DM infection levels significantly and increase grain yield. Downy mildew infection decreased by between 58 and 100% while grain yield increases ranged from 10 to 98% for the 2‐4 cycles of selection relative to the C0 (original). Selection increased grain yield with acceptable changes in plant height while maintaining maturity in disease‐free environments.     

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.     

Yield response to honeycomb selection in maize

Summary This research evaluated the response to three cycles of mass honeycomb selection based on the grain yields of widely-spaced plants from the F₂ generation of a maize (Zea mays L.) single cross hybrid. Frequency distributions of individual plant yields in the selection plots showed strong positive skewness. A significant linear yield increase of 11.23% per cycle compared to the check was obtained. Correlated responses were the significant reduction in days to mid-silking, increases in ear and plant heights and in prolificacy. The selected population was similar to the original hybrid in days to mid-silking, ear and plant heights, but had significantly more ears per plant. It is concluded that mass honeycomb selection at wide spacings was effective in improving grain yield and prolificacy but may produce correlated responses in some other agronomic traits similar to those obtained with other selection schemes.     

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.     

Mass selection of wheat for grain filling without transient photosynthesis

Summary Post-anthesis chemical desiccation of wheat (Triticum aestivum L.) plants in the field eliminates transtent photosynthesis by killing all green tissues, thus revealing the plant's capacity for grain filling from stored stem reserves, as the case is for post-anthesis stress such as drought or leaf diseases. This study was conducted to investigate whether mass selection for large kernels under chemical desiccation would lead to the improve ment of grain filling in the absence of transient photosynthesis.Six crosses of common spring wheat were subjected to three cycles of mass selection from F₂ through F₁ when selection was performed for large kernels by sieving grains from plants that were erther chemically desiccated after anthesis, or not (controls). The resulting 36 bulks (six crosses by three selection cycles by two selection environments) were compared with their respective F₂ base populations, when tested with and without chemical desiccation.Selection for large kernels under potential conditions (without chemical desiccation) did not improve kernel weight under potnetial conditions, evidently because these materials were lacking in genetic variation for kernel weight under potential conditions. In four of the crosses, 3rd cycle selection for large kernels under potential conditions decreased kernel weight under chemical desiccation. On the other hand, selection for large kernels under chemical desiccation was effective in improving kernel weight and test weight under chemical desiccation, depending on the cross and the selection cycle, with no genetic shift in mean days to heading or mean plant height. Selection for large kernels under chemical desiccation was also effective in some cases in increasing kernel weight under potential conditions. The results are interpreted to show that selection under potential conditions and under chemical desiccation operate on two different sources for grain filling, namely transient photosynthesis and stem reserve utilization, respectively. In order to expose genetic variability for stem reserve utilization to selection pressure, transient photosynthesis must be eliminated, as done by chemical desiccation in this study.     

Direct and Correlated Responses to Four Cycles of Divergent Selection for Heading Date in Barley*

In order to investigate the direct and correlated responses to selection for heading date in barley, two F₂ populations were subjected to four cycles of divergent selection. For each cross, eight populations selected for earliness (from 1E to 4E) and for lateness, (from 1L to 4L) and four unselected populations (1, 2, 3 and 4 SSD) were obtained and than compared together with parental lines. Selection caused a change greater than one day/cycle in Doth directions, and the cumulative responses were -5.4 and -5,9 days in the direction of earliness, 7.7 and 7,1 days in the opposite direction. Such results suggest that in the two base populations the heading, date is under the control of a polygenic system with prevailing additive effects and a slight directional dominance toward earliness. Early populations showed a reduction of ear moisture and an increase of plant height, ear extrusion, kernel weight and harvest index, whereas late populations showed the opposite changes. Grain yield was linearly reduced by selection for lateness, whereas no substantial change was evidenced in the opposite direction, so that 4E significantly out-yielded 4L in both crosses.     

Estimates of additive and dominance effects for Fusarium head blight resistance of winter triticale

The most effective strategy to control Fusarium head blight (FHB), a devastating disease of small‐grain cereals, is breeding resistant cultivars. This resistance study of F₁ crosses, F₂ and backcross generations of triticale estimates heterosis, general and specific combining ability (GCA, SCA), additive and dominance effects and compares parents with segregating generations. The genetic material consisted of 10 parents with their 45 F₁ crosses and of six parents with their 15 F₂ progeny and backcrosses to each parent. Genotypes were grown in various environments and artificially inoculated with an aggressive isolate of F. culmorum. FHB was assessed, by visual rating, as the mean of four to five individual ratings of disease development. Heterosis for FHB was of little importance. The correlation between the FHB rating of F₁ crosses and their mid‐parent performance was close. GCA was the predominant source of variation, although the significance of the SCA variance also implied non‐additive allelic interaction. The preponderance of additive gene effects is encouraging for increasing resistance by a recurrent selection programme. The relationship between the GCA effect of a parent and its per se performance was close, which gives the possibility of predicting FHB resistance in F₁ crosses. Additive effects were predominant in the F₂ progeny and also in the backcrosses. Transgressive segregants could not be detected. Searching for them should be postponed to the F₃ or later generations.     

Responses to selection and changes in combining ability after three cycles of a modified reciprocal recurrent selection in maize

This research reports responses to selection and changes in general (GCA) and specific (SCA) combining abilities after three cycles of a modified reciprocal recurrent selection procedure (MRRS) in EPB-4 and EPB-5 maize populations. In the MRRS procedure a cycle can be completed in 1 or 2 years depending on the availability of winter breeding nurseries. The original and the three selection cycles of the two populations per se (eight entries) and the partial diallel developed from the crosses between them (16 entries) were evaluated in six environments. Realized response to selection on the population cross was 7.25% cycle⁻¹ for grain yield, −13.63% cycle⁻¹ for plant lodging, and 11.93% cyle⁻¹ for prolificacy, whereas plant and ear heights remained unchanged. GCA estimates increased with selection cycles for both populations for grain yield and prolificacy, and decreased for plant lodging, indicating that the frequency of favorable alleles with additive effects for these traits increased with the MRRS cycles in both populations. SCA estimates increased for grain yield and prolificacy indicating that the frequency of favorable complementary alleles at loci with non-additive effects in the reciprocal populations increased with the MRRS cycles. For grain yield, SCA effects increased more than GCA effects with selection cycles, indicating that MRRS exploited more the non-additive effects than the additive effects for the improvement of this trait. The overall results showed that the MRRS procedure was highly effective in improving the population cross, exploiting both GCA and SCA effects.     

A comparison of yield and yield component selection in winter wheat

Summary Three populations of winter wheat were formed by crossing Avrora to Sage, TAM W-101, and Danne. Approximately 10% of the F₂ plants from these crosses were selected for high and low levels of number of tillers per plant, number of kernels per spike, 1000-kernel weight, and grain yield. Forty-eight solid seeded F₃ lines obtained from the selected F₂ plants were then selected for high and low expressions of yield components and grain yield. Realized heritabilities were estimated. Indirect responses of yield to yield component selection and direct response to selection for grain yield were measured. Heritabilities were low for tiller number, number of kernels per spike and kernel weights but were high or intermediate for grain yield when selection occurred in the F₂ generation. When selection was practiced in the F₃ generation, heritabilities for tiller number and yield were low, but were intermediate to high for number of kernels per spike and kernel weight and high heritabilities were found for kernel weight. Selection for kernel weight often increased grain yield; however, direct selection for grain yields was usually as effective.Journal article no. J-4488 of the Oklahoma Agri. Exp. Stn., Stillwater, Oklahoma 74074.     

Selection in Early Generations of Dried Peas, Pisurm sativum L.

Seed production in peas is limited by early and severe lodging and by a low multiplication rate and pea breeders have tried to overcome such disadvantages by growing plants in early generations widely-spaced against wire netting. Progenies of four pea crosses were each tested over a period of tour years to investigate whether single plant selection for quantitative characters, in particular grain yield, is efficient in such circumstances in the Is generation. Efficiency of selection was also studied for yield and yield related characters in the F₃ based on f₂ derived single plant progenies grown in unreplicated microplots. Early yield selection in the F₂ as well as in the F₃ proved to be of poor efficiency. Heritability values of all recorded traits were only slightly higher in the F₃ than in the F₂. Efficiency of yield selection could not be increased significantly in these early generations by indirect selection, either for single traits or for index traits. Possible reasons are discussed for the low efficiency of direct and indirect selection for grain yield in early generations and proposals are made for its improvement and for alternative selection strategies.     

Genetic progress after four cycles of recurrent selection for yield and grain traits in common bean

The objective of this study was to evaluate the genetic progress after four cycles of recurrent selection in common bean. The base segregating population was obtained from 10 parents, and derived the S_0:1 and S_0:2 families that were evaluated. The S_0:3 families with higher grain yield and grain color, like the standard carioca were selected, and were intercrossed to generate the population of the following cycle. This process was repeated for four cycles. The best families were evaluated in each cycle by many generations and locations, and the five best lines of each cycle were identified. The 20 lines thus obtained were evaluated in two growing seasons, sown in July and November 2002. The grain yield (kg/ha) and grain type (scale of scores) were evaluated. Genetic progress was confirmed for both traits. The mean annual gain with selection for the grain type was 10.5% and 5.7% for grain yield, with no evidence of variability reduction in the population. These results show that recurrent selection is a good alternative for improving common bean quantitative traits.     

A recurrent selection programme for grain yield in winter barley

Summary The effect of a recurrent selection procedure was evaluated in a winter barley (Hordeum vulgare L.) population. Cycle zero (C₀) was initiated by crossing six high yielding winter barley cultivars with the short straw cv Onice. The F₁'s were crossed according to a diallel scheme without reciprocals. A total of 750 S₀ plants were derived and evaluated; 329 S₀ plants were selected and their progenies (S₁ lines) tested. Fifteen S₁ lines were chosen and used as parents of cycle 1 (C₁), by producing 105 F₁ hybrids which simulated a random mating offspring. One hundred and three randomly chosen S₁ lines belonging to C₀, and 103 S₁ lines belonging to C₁, were evaluated at two locations.For grain yield a significant difference between cycles was observed. From C₀ to C₁ the grain yield increased with 307 g/m². This increase was due to a higher number of seeds per m². For plant height, heading date and 100-kernel weight no differences between cycles were observed.The positive results obtained in this study indicate the potential usefulness of recurrent selection for developing parents or lines superior for grain yield, with little change in other important agronomic traits.     

Evaluation of Pedigree and Single Seed Descent Selection Methods for Cultivar Development in Cowpea (Vigna unguiculata L. Walp)

Three crosses of cowpea (Vigna unguiculata [L.] Walp) were used to assess the effectiveness of two selection procedures in cowpea. The selection procedures were pedigree selection (PD) and single seed descent selection (SSD). Lines developed through each procedure were yield tested and selected at F₆ and F₇ for PD and SSD, respectively. The selected lines for both procedures were compared at F₈ for grain yield per plant (YLD), number of pods per plant (NPO), and pod development period (PDP), in two locations. Lines developed through both procedures differed more in their population means than in the performance of their top 10% or highest yielding lines. There was no consistent relationship between magnitude of genetic variability and percentage of superior lines obtained from each procedure. Both procedures have been effective in producing superior genotypes for yield and number of pods. The PD method produced superior transgressive segregants in two out of the three crosses for PDP. SSD allowed a more rapid generation than PD.     

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.     

Inheritance pattern of mungbean yellow mosaic disease resistance and gene action for different traits in mungbean (Vigna radiata (L.) Wilczek) under protected and unprotected field conditions

Understanding of mungbean yellow mosaic disease resistance is crucial to develop resistant varieties to combat worldwide threat of the disease epidemics. This study was aimed to determine nature of disease resistance in terms of number of genes governing it and genetics of related traits. Experimental site was located on 31.43°N and 73.06°E with an elevation of 186 m, and evaluation trials were conducted during spring season due to high disease epidemics in this season. Two contrasting genotypes, that is, NM 6‐68‐2 (resistant) and NM 1‐32‐1 (highly susceptible), were crossed to raise six populations, that is, P₁, P₂, F₁, F₂, BC₁ and BC₂ for evaluation under protected and unprotected field conditions using chi‐square test and generation mean and variance analysis. It was discovered that disease resistance was governed by two major genes with additive effects. Disease resistance can easily be incorporated through backcrossing and direct selection following hybridization. Direct selection should be practised at earlier generation for plant height, chlorophyll contents and number of seeds per pod due to preponderance of additive effects whereas at later generations for seed yield per plant, 100 seed weight, harvest index, number of pods per plant and pod length due to involvement of duplicate epistasis.