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.     

The Relationship Between Progeny Seed Yield in Drilled Plots and Maternal Spaced-Plant Traits in Perennial Ryegrass (Lolium perenne L.)

Seed production in perennial ryegrass, Lolium perenne L., is low. Early selection for high seed yield carried out in spaced plants would be valuable for breeders, but only if this potential is also expressed in derived progenies sown later in drilled plots. Therefore, spaced-plant traits assessed on clonal ramets of 31 genotypes in each of two perennial ryegrass varieties in four environments were related to seed production characteristics in drilled plots of open-pollinated progenies of these plants. Significant differences were found among these half-sib families for seed yield, thousand-grain weight, spring performance and earliness. The number of spikelets per ear in the maternal plants was negatively correlated with the seed yield of the progenies, explaining 17 % of the variation for seed yield in the cv. ‘Barenza’ and 14 % in the cv. ‘Wendy’. Multiple regression analyses revealed that no combinations of plant traits consistently explained a major portion of the variation for seed yield of the progenies. The results indicate that in these varieties, spaced-plant data are of limited value in predicting seed production. Direct selection for seed yield in drills of progenies in later stages of the breeding programme remains the best method for obtaining varieties with sufficient seed production.     

Associations of some characters with seed yield in chickpea collections

Summary Three thousand two hundred and sixty-seven kabuli chickpea (Cicer arietinum L.) germplasm accessions were grown during the spring season of 1980 at Tel Hadya, the main research station of ICARDA, Syria to determine the components of seed yield. Observations were recorded on seed yield and 14 other characters. Correlation and path coefficient analyses were done to find out associations among characters and to assess the direct and indirect contribution of each character to seed yield.Large variation was observed for all the characters studied except days to flowering, days to maturity and protein content. Correlation and path coefficient analyses showed that biological yield and harvest index were the major direct contributors to seed yield. The 100-seed weight, plant height, days to flowering and maturity, canopy width, and protein content contributed to seed yield mainly through indirect effect via biological yield and harvest index. The 100-seed weight and seed yield were major contributors to biological yield. Major contributor to protein content was days to maturity. Results indicated that selection for high biological yield and harvest index would lead to high seed yield; and selection for large seed size would lead to high biological yield. Therefore, these characters should receive the highest priority in selecting high yielding plants in chickpea breeding.     

Prediction of indirect selection for seed and forage yield of lucerne based on evaluation under spaced planting

Phenotypic selection under spaced planting is frequently used in forage species. This study aimed to compare the predicted efficiency of direct selection for lucerne seed or forage yield under dense planting with indirect selection based on the same traits or seed yield components evaluated under spaced planting. Sixteen genotypes randomly chosen from a representative sample of locally adapted germplasm were grown for two years as individual clones spaced at 75 cm (density =1.78 plants/m²) and in dense plots formed by one row of four clones spaced at 10 cm (density = 50 plants/m²) using a randomized complete block design with three replications. Indirect selection based on seed yield under spaced planting was just 19% less efficient than direct selection, owing to moderate genetic correlation between plant densities (r_g = 0.66) and somewhat higher broad‐sense heritability under spaced planting than under dense planting. The relative efficiency of indirect selection for seed yield in density based on individual seed yield components under spaced planting ranged from modest to very low and was always below 45%. The efficiency of indirect selection for dry matter yield based on yield response under spaced planting was moderate for total yield (64%) and very low for second‐year yield (32%) relative to direct selection.     

Improved seed yield in perennial ryegrass (Lolium perenne L.) from two generations of phenotypic selection

Two generations of recurrent phenotypic selection for seed yield per plant under controlled pollination were carried out to combine improved seed yield with improved forage grass performance in perennial ryegrass (Lolium perenne L.). Selected and unselected varieties (AberDartand AberElan respectively) and control varieties were grown for seed in pots in a glasshouse experiment and in two field plot experiments over 5 harvest years. Seed yield components and seed yield were measured. Significant improvements in seed yield of the selected AberDart over the unselected AberElan were observed both in individual plants grown in the glasshouse and in field plots. Detailed observations of the individual components of seed yield showed that the increased seed yield of AberDart can be attributed to a higher proportion of ovules forming seeds (% seedset), greater seed number per tiller and more reproductive tillers per plant. This improvement in seed yield of AberDart was also observed when it was grown in field plots with AberElan, experimental selection lines and commercial varieties. AberDart produced more seeds per tiller than the other commercial varieties and selection lines studied. The implication of this approach for the breeding of perennial ryegrass varieties that combine good agronomic performance and acceptable seed yields is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Recurrent selection for yield in Glycine max using genetic male-sterility

Summary Recurrent selection, using genetic male sterility to facilitate intermating in Glycine max, was evaluated when selecting among individual space-planted S₀ soybean plants for three yield-related traits: 1) seed yield per se (YLD), 2) apparent harvest index (AHI), and 3) a selection index in which seed yield was regressed on maturity (REG). The original intermating population was a combination of the cultivar Century and two F₂ populations segregating for male sterility. The selection intensity through three cycles of selection was 20%. The YLD population increased in mean yield by 2.8±2.2 g plant^-1 cycle^-1 buf shifted significantly toward late maturity by 3.9±0.6 days cycle^-1. The AHI population decreased in yield by 4.4±2.6 g cycle^-1 while maturity shifted slightly earlier; there was very little change in AHI. The REG population increased in yield by 5.7±2.4 g plant^-1 cycle^-1 while shifting toward later maturity by only 0.8±0.5 day cycle^-1. We concluded that selection among S₀ plants in these populations, using the selection index of yield regressed on maturity, was affective in increasing single plant yields without causing a significant shift in maturity. Selecting for yield per se caused an unacceptable shift toward late maturity and selection for AHI was ineffective for increasing yield or AHI.     

Genetics of Seed Yield and its Components in Common Bean (Phaseolus vulgaris L.) of Middle-American Origin

Genetic variance, heritability, and expected response from selection arc useful in devising alternative methods and criteria of: selection. The objectives of this study were to estimate these for seed yield and its components from 200 F₂: populations involving 80 cultivars and lines of mostly small-seeded dry bush bean (Phaseolus vulgaris L.) of habits growth I, II, and III of Middle-American origin. All cultivars and lines were crossed in eight sets of ten parents each in a Design II mating system. The F₂ populations, without parents, were evaluated in the field in a replicates-in-sets design at two locations in Colombia in 1983. Estimates of additive genetic variance were significant for yield, pods/m², seeds/pod, and seed weight. Interaction with environments was also significant. Values for nonadditive genetic variance were not significant for either yield or yield components. The estimates of narrow sense heritability, based on the F₂ population mean and unbiased by genotype x environment interaction, were 0.21 ± 0.13 for yield. 20 ± 0.13 for pods/m², 0.57 ± 0.13 for seeds/pod, and 0.74 ± 0.15 for seed weight. The expected direct response from selection of the top 20 % of F₂ populations for yield per se would result in a 4.30 % increase in yield with a correlated response of 0.21 % in seed weight. In contrast, the expected gain from direct selection for seed weight would result in a 11.76 % increase in seed weight with a, correlated gain of 0.28 % for yield. Direct selection for pods/m² would decrease yield, seeds/pod and seed weight, while direct selection for seeds/pod would reduce pods/m² and seed weight but increase seed yield by 0.37 %. Data on yield from replicated trials in the early segregating generations could be utilized for identification and selection of promising crosses and families or lines with crosses for dry bean yield improvement.     

Molecular markers for yield components in Brassica juncea – do these assist in breeding for high seed yield?

A population of 112 F₁-derived doubled haploid lines was produced from a reciprocal cross of Brassica juncea. The parents differed for seed quality, seed color and many agronomic traits. A detailed RFLP linkage map of this population, comprising 316 loci, had been constructed, and was used to map quantitative trait loci (QTL) for seed yield and yield components, viz. siliqua length, number of seeds per siliqua, number of siliques per main raceme and 1000-seed weight. Stable and significant QTLs were identified for all these yield components except seed yield. For yield components, a selection index based on combined phenotypic and molecular data (QTL effects) could double up the efficiency of selection compared to the expected genetic advance by phenotypic selection. Selection indices for high seed yield, based on the phenotypic data of yield and yield components, could only improve the efficiency of selection by 4% of the genetic advance that can be expected from direct phenotypic selection for yield alone. Inclusion of molecular data together with the phenotypic data of yield components in the selection indices did not improve the efficiency of selection for higher seed yield. This is probably due to often negative relationships among the yield components. Most of the QTLs for yield components were compensating each other, probably due to linkage, pleiotropy or developmentally induced relationships among them. The breeding strategy for B. juncea and challenges to marker assisted selection are discussed.     

Recurrent selection for maturity and percent seed protein in Glycine max based on S0 plant evaluations

Summary Progress was evaluated after four cycles of recurrent selection among S₀ plants of Glycine max (L.) Merr. in which selection was either for maturity (MAT) or seed protein (PRO). The two populations, MAT and PRO, were developed from an initial population that was a combination of a line with 48.4% seed protein and two F₂ populations segregating for male sterility. Intermating was facilitated by genetic male-sterility and the selection intensity was 20% in each cycle of the two populations. Selection for early maturity advanced the average maturity a significant 2.7±0.34 days cycle^–1 and reduced seed yield a non-significant 9.1±2.95 g plant^–1 cycle^–1. Selection increased mean seed protein a significant 0.8±0.15 percentage points cycle^–1 and decreased percent seed oil a non-significant 0.5±0.17 percentage points cycle^–1. Correlation coefficients between seed protein and seed yield varied from 0.18 to –0.21 in the four cycles indicating plants with favorable combinations of seed yield and seed protein could be identified. Selection in these two populations would be effective for early maturity and for increased seed protein.     

Geneticanalysis of drought stress response in rapeseed (Brassica campestris and B. Napus). II. Yield improvement and the application of selection indices

Summary The expected improvement in grain yield if selection was made for plant characters measured between flowering and maturity was evaluated in populations of rapeseed (Brassica campestris and B. napus) grown in a droughted environment. Drought was commenced at flowering in each species and measurements were made on plants at the commencement of the drought stress, during the stress treatment and at crop maturity.Substantial genetic and phenotypic variation was observed in yield as well as the different morphophysiological determinants of yield. In B. campestris no single parameter was found to be a suitable alternative selection criterion to yield since the correlated responses in yield if selection was for another character was lower than if selection was for yield alone. By the use of a selection index however, joint selection for yield, as well as harvest index, 1000 seed weight and seeds per pod, was expected to be 20% more effective than direct selection for yield under drought. In the B. napus population direct selection for flowering time or for harvest index was predicted to result in a genetic advance in yield equal to or greater than that obtained by direct selection for yield, whereas joint selection for flowering time and yield should result in a 16% greater yield increase. Selection for increased yield in these populations grown in droughted conditions is discussed.     

Effect of plant density on selection for seed yield in two population types of Phaseolus vulgaris L.

Summary Pedigree selection for seed yield, using early generation yield tests, was practiced from the F₂ to F₇ in two populations of common bean (Phaseolus vulgaris L.) in three plant densities: 66, 133, and 266 thousand plants ha^-1 at CIAT-Palmira, Colombia. The six highest yielding lines selected from each plant-density, along with 13 parents, were evaluated in a 7×7 lattice design with three replications at the three densities utilized for selection, and at 399,000 plants ha^-1 in 1988 and 1989.Based on the mean performance of selected lines and the mean of the parents, selection for seed yield was effective in all densities in both populations. However, none of the lines selected from the population within the race Mesoamerica (TC 4673) significantly outyielded their best parent under any plant density. The highest yielding lines selected from the interracial population (TR 4635) outyielded their best parent irrespective of the plant-density used for selection. The highest yielding line originated from the highest density used for selection. Low density was neither good for selection nor for evaluation and identification of high-yielding cultivars of common bean. There was no significant difference between the mean yield of lines selected at the intermediate and high population densities. The effects of plant density, year, and their interactions were significant for seed yield.     

Improvement of large-seeded common bean cultivars under sustainable cropping systems in Spain

Approaches are needed to broaden the genetic base and improve earliness and yield potential of large-seeded beans under sustainable cropping systems. The objective of this research was to develop adapted dwarf bean populations having a commercial seed quality and yield suitable for the production in the South of Europe. The original base populations were produced from crosses between genotypes within each Mesoamerica, Nueva Granada and Peru races, and between Peru and Nueva Granada, and Mesoamerica and Nueva Granada races. Visual mass selection for plant performance was practised in the F₂ and F₃ generations. In the F₄ and F₅, single plants were harvested under two cropping systems (sole cropping and intercropping with maize). From F₄, selection was based on precocity, combined with seed yield and seed commercial type. The F_4:7 selected lines from each original population were compared with their parents and five checks at four environments and two cropping systems. Differences among environments, populations, parents and checks were observed for all traits. Under intercropping with maize, there was a 50% reduction in seed yield. Yield of parents and checks belonging to Andean South American races, intraracial (Nueva Granada × Nueva Granada) and interracial (Nueva Granada × Peru) populations, was higher than that of those of Middle American origin. Intraracial crosses within large-seeded Andean South American (Peru race) and Middle American gene pools (Mesoamerica race) did not produce lines yielding more than the highest yielding parent. Only two large-seeded lines selected from crosses between small- and large-seeded gene pools out-yielded the best parent and check cultivar.     

Yield components,harvest index and plant type in relation to yield differences in field pea genotypes

Summary The effectiveness of yield components, harvest index and morphological characteristics as selection criteria among four field pea (Pisum sativum L.) genotypes was examined. Genotypes were grown at a wide range of plant populations (9 to 400 plants m^-2) to maximise environmental diversity.Both biological and seed yields approximately doubled from 9 to 100 plants m^-2. This response flattened from 100 to 400 plants m^-2. Differences among the genotypes were found only at 225 and 400 plants m^-2. Analysis of the yield components highlighted the plasticity and large genotype by environment interactions of field peas. The numbers of pods per plant and peas per pod were maximised when each genotype was grown as spaced plants, but the low plant numbers meant seed yields per unit area were at their lowest.Genotypic differences for plant harvest index (PHI) were also only found at 225 and 400 plants m^-2. Broad sense heritability estimates indicated that direct selection for PHI was not feasible. The inference from the yield component and PHI results was that alternative selection criteria such as physiological or morphological characteristics may be necessary for improved yield potential. Classification of each genotype indicated that low seedling vigour may be a positive attribute for crop plants of semi-leafless and conventionally leafed field peas. Selection based on any single plant attribute is unlikely to lead to dramatic improvements in the yield potential of field peas. Selection should be based on plant characteristics rather than on differences in yield components.     

Heritability estimates of spaced-plant traits in three perennial ryegrass (Lolium perenne L.) cultivars

Summary Seed yield in perennial ryegrass is low and unpredictable. Spaced-plant traits suitable for indirect selection for total seed yield in drilled plots would be very useful. The objectives of this investigation were to evaluate genetic variation for seed yield components and other traits among clones from three perennial ryegrass cultivars differing in seed yield and their open-pollinated progenies. Per cultivar, a random set of 50 genotypes was cloned and on each genotype seed was generated by open pollination. Clonal ramets of the parents were observed for 17 traits in 1986 at two locations. In 1987 and 1988, parents and progenies were observed as single plants in a randomized complete block design with two replications. There was little cultivar-environment interaction for most traits. The parents differed significantly for almost all traits. Half-sib (HS) families differed for only three to five traits. Broad-sense heritabilities (h² _b), based on variance components of the parents, were moderate to high; earliness had the highest h_b ². Narrow-sense heritabilities (h² _n), based on variance components among HS-families, were low to moderate and mostly not significant; for most traits h² _n estimates varied between years and cultivars. Flag leaf width and date of first anthesis showed the highest h² _n. Narrow-sense heritability estimates from parent-offspring regressions (h² _nPO) ranged from non-significant to high, depending on year and cultivar; they were generally higher than the corresponding h² _n estimates. Generally, h² _nPO was highest for earliness, flag leaf width, ear length and the number of spikelets per ear. Breeding methods that capitalize on additive genetic variance, such as mass selection, should result in improvement for these traits.     

Selection for Increased Seed Nitrogen Accumulation in Common Bean: Implications for Improving Dinitrogen Fixation and Seed Yield

Estimates of N₂ fixation in segregating populations of bean plants based on ¹⁵N-isotope methods are technically demanding and expensive; therefore, indirect measures based on related traits including total seed N were used to select for improved N₂ fixation and yield. In 1985, six populations of F₂-derived F₃ families resulting from six parental lines crossed to a common tester were grown in field trials on a low-N soil. In 1986, 25 selected half-sib families and two populations of full-sib F₄ families were grown under similar conditions. Parents and a non-nodulating soybean line were included both years. Narrow sense (H_NS) heritability estimates based on parent-offspring regressions ranged from 0.57 for total seed N to 0.39 for shoot biomass in one population, but were near zero for all measured traits in a second population. Among the criteria used to identify parental lines with superior potential for producing progenies with high total seed N, testcross population means combined with estimates of realized heritability were the most reliable. Selection of the best F₃ families for total seed N resulted in F₄ families with increased total seed N and higher seed yields, while seed protein percentage was unchanged. When plants are grown on low-N soils, selection for total seed N offers a useful alternative to selection for increased N₂ fixation based on ¹⁵N-isotopic or total-N-difference method.     

Response to selective pressure in early generation progenies of upland cotton (Gossypium hirsutum L.)

Summary Combining high fiber strength with high yield in upland cotton (Gossipium hirsutum L.) was been difficult. The cross combination, 69–120 × 6M-10, was chosen for this study because of the divergence of parents for fiber strength, seeds per boll and crop maturity. Forty F₂ plants were selected in 1974 solely on the basis of visual yield and 20 plants were randomly chosen to serve as a bulk check. The F₃ progenies, the bulk check, the parents and a commercial check were tested in 1975. From these results, five groups were established, consisting of four (10%) progenies each to represent high seed/boll, low seeds/boll, high fiber strength, low fiber strength and high yield in the 1976 test of F₄ progenies. This experiment was conducted to (a) compare the contributions of yield components to lint yield among groups of early generation progenies (F₃ and F₄) and (b) determine the phenotypic correlations between yield and quality attributes. The yield components, bolls/m², seeds/boll, fibers/seed, mean fiber length and micronaire, were included as well as fiber strength.No significant lint yield differences were found among the five groups. Progenies within groups did, however, differ in lint yield. The low and high seeds boll groups gave similar yield and fiber quality results. The low and high fiber strength groups gave similar yield but longer fiber was obtained with the high strength group.Stepwise regression analyses estimates show that the number of bolls produced per unit area, although the major contributor to lint yield for all groups, contributed only 66.9% of the total variation in the progeny group selected for high yield, with fibers/seed, mean fiber length, micronaire and seeds/boll accounting for 13.2%, 8.1%, 6.0%, and 5.8%, respectively. Correlations between the various yield components and fiber strength were low and mainly insignificant, indicating that minor alterations might be made in later generations without serious consequences.Potential gains from second-stage selective pressure upon yield-per-boll components following initial selection for lint yield are discussed.     

Selection for Seed Yield Production of Egyptian Clover (Trifolium alexandrinum L.) C. V. Fahl

Modified mass and family selection for seed yield production of berseem clover (Trifolium alexandrinum L.) c.v. ‘Fahl’ were applied for two generations. Two hundred plants (5 % intensity of selection) were selected for seed yield in the first season, 1985/86. In the second season, 1986/87, selection between and within half-sib families was practiced. In addition, equal parts of seeds from each of the 200 selected plants were bulked to form the C₁ modified mass selection; after establishing, the same procedure was adopted to form the C₂ generation. The cycles 1 and 2 of half-sib families and modified mass selection along with the base population family were evaluated for forage and seed yields. The realized gains from modified mass selection were 6.03 and 9.51 % for fresh forage yield, 5.57 and 10.86 % for protein yield and 13.23 and 16.19 % for seed yield in cycles 1 and 2, respectively, over the base population. The realized gain from family selection in cycle 2 as a percentage of the base population mean amounted to 11.32, 13.35, 17.47 and 3.15% for forage, protein, and seed yield and seed index, respectively. The broad sense heritability, as estimated from the variance components was 89.63, 63.03, and 76.67 % for dry forage, seed yield and seed index, respectively. Although, all these five traits (fresh, dry, protein, and seed yield and seed index) had positive correlation with each other, weak correlations were found between seed yield and forage yields. Furthermore, close associations were found among forage yield 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.     

Indirect selection for persistence and seed yield in large-leaved white clover

High persistence and seed yield are important targets in white clover breeding but their assessment is expensive. The efficiency of indirect selection for these traits performed on characters recorded during a medium-term dry matter (DM) yield evaluation was verified for large leaved populations in separate experiments for medium-term (three-year) and long-term (fourth and fifth year) DM yield (Trial 1) and seed yield (Trial 2). The best results were provided by stolon density observed at the end of the third cropping year for prediction of long-term yield, and by the product of number of heads times number of florets per head assessed prior to mowing in Trial 1 for prediction of seed yield in Trial 2. Indirect selection was about as efficient as direct selection in these cases. Predictions of persistence from three-year or third year DM yield, and seed yield from single seed yield components in Trial 1 were less accurate.     

Selection for resistance to Septoria nodorum in wheat

Summary A population of 572 F₂ derived F₃ lines from six crosses were used to estimate parameters relevant to selection for resistance to Septoria nodorum of wheat. Lines were grown in disease free (fungicide sprayed) and inoculated microplots in 2 replications of a split-plot design in a single environment in 1977. Average yield reduction due to disease was approximately 50%; this was associated with an average septoria score of 50% on the flag leaf, an average septoria score of 42% on the head, and a reduction of 37% in seed weight. Low S. nodorum scores were correlated with late heading date, tall plant height, high grain yield, and high seed weight in diseased plots, and high seed weight % (seed weight in diseased plots expressed as a percentage of seed weight in fungicide sprayed plots).Restricted selection indexes were used to study the relative contributions of disease escape, true resistance, and tolerance to variability in grain yield in diseased plots, seed weight in diseased plots, and seed weight %. True resistance appeared to be the most important factor causing variation in grain yield in diseased plots and seed weight %. Tolerance and escape seemed to be more important for seed weight in diseased plots.Heritabilities of S. nodorum scores on the flag leaf and head were 63% and 52%, respectively. Leaf and head scores could be used most effectively as selection criteria to upgrade resistance in a population before harvest.Selection for high seed weight % slightly reduced yields in disease free plots, although yield in diseased plots and seed weight in diseased plots were increased. However, selection for increased yield or increased seed weight in diseased plots improved yield in disease free plots. It is suggested that direct selection for yield or seed weight in diseased plots is likely to achieve more desirable goals than selection for seed weight %.