Accelerated recurrent selection

In Accelerated Recurrent Selection (ARS) schemes, selection is based on the predicted performance of progeny families, estimated as the average of the parental families. These schemes can halve cycle time compared to simple recurrent selection methods. They also have a lower requirement for evaluation of families in yield trials, although they do require more seed production. ARS therefore provides options in cycle time, effective population size, response to selection and cost which have not been available before. Example schemes are compared by computer simulation with truncation selection and with optimal family selection, where contributions of families to the next generation are optimised to give the maximum response to selection at a specified effective population size. In many circumstances, ARS schemes compare favourably. Difficulties in combining estimates of selection intensity and of effective population size when comparing the merits of different breeding schemes are discussed. It is suggested that unless one is interested in response to selection over periods greater than 50 years, the weight given to effective population size in ranking different schemes should be small.     

Reciprocal recurrent selection for multitrait indices in maize

Summary Reciprocal recurrent selection was carried out with two populations of maize (Zea mays L.) having good combining ability. We selected for higher grain yield, early maturity, shorter plant height and lodging resistance. Two cycles were completed in two years (four seasons), by resorting to late planting of S1 lines for recombination in the main season in which top-cross families were assessed. Top-crosses and selfings were made in the off-season. The original and improved versions of the populations and their crosses were evaluated in multilocation trials. The superiority of the population hybrid was 10.3% for grain yield, 5.5% for plant height and 1.8 days to silk. The improved hybrid had delayed leaf senescence and better resistance to lodging and post-flowering stalk rots. Among the yield components, only ear girth showed improvement. Syn 2 of the improved population cross showed a yield reduction of 6.2% in comparison to Syn 1. Intrapopulation gains were not significant except for lodging resistance in one population.     

Correlated response to recurrent selection for groat-oil content in oats

Summary One hundred random oat (Avena sativa L.) lines from a base (C₀) and each of three populations (C₁, C₂, and C₃) improved for groat (caryopsis) oil content by phenotypic recurrent selection were evaluated for correlated changes in several unselected agronomic traits. In addition, the parents of the base population and four check varieties were evaluated for the same traits. Phenotypic recurrent selection for high groat-oil content resulted in no significant correlated response in mean expression of any trait. Mean grain yield, biomass, groat yield, and harvest index of the improved populations were equal or superior to the mean of the parents and, with the exception of harvest index, equivalent to the mean of the check varieties. Mean test weight and seed weight of all populations were lower than for parents or check varieties. Selection for high groat-oil content caused a decline in genotypic variance for test weight and groat fraction, but reductions in genotypic variance for heading date and plant height may have resulted from culling for good agronomic type. Broad-sense heritability remained moderate to high for all traits except groat fraction. Phenotypic and genotypic correlation coefficients revealed negative, though mostly nonsignificant, relationships between groat-oil content and several traits, which may reflect a purported bioenergetic limitation to increasing groat-oil content in oats. Oil yield, however, was positively correlated with grain and groat yield, groat fraction, biomass, and harvest index. Results suggest that development of high-oil oat cultivars with current levels of production traits via phenotypic recurrent selection is possible.Journal Paper no. J-13038 of the Iowa Agric. and Home Econ. Exp. Stn., Ames, IA 50011. Project 2447.     

Genetic diversity among populations and breeding lines from recurrent selection in Brassica napus as revealed by RAPD markers

Recurrent selection facilitated by dominant male sterility has been conducted to broaden the genetic basis for cultivar development in Brassica napus. This study aimed to evaluate the genetic variation in four base populations (C0‐C3) and breeding lines from two of the populations produced during recurrent selection by random amplified polymorphic DNA (Rapd) markers. Genetic variation in four populations declined gradually with the advance of selection cycles as measured by expected genetic heterozygosity (from 0.2058 in C0 to 0.1536 in C3) but the decline was not statistically significant. When compared with the average genetic distances for 21 germplasm collections with wide geographical and genetic origins (0.4712) and seven breeding lines from pedigree selection (0.2059), seven breeding lines selected from the C1 population and 11 from the C3 population had a larger average genetic distance (0.5339 and 0.5486, respectively). Clustering analysis indicated that the lines from recurrent selection had a much lower genetic similarity than lines from pedigree selection. Our results suggest that base populations derived from recurrent selection could provide a wider genetic variation for selection of breeding lines with more broad genetic bases.     

Changes in genetic diversity during seven cycles of recurrent selection for grainyield in oat, Avena sativa L.

Variability for desirable alleles within elite breeding populations is a requirement for long-term genetic improvement. Changes in genetic diversity were evaluated in a recurrent selection programme for enhanced grain yield in oat, Avena sativa L., using coefficient of parentage (r_p) and restriction fragment length polymorphism (RFLP) based genetic similarity (S) estimates. Both r_p and s increased during seven cycles of recurrent selection, indicating a reduction in the level of genetic diversity within the population. The relationship between genetic diversity and agronomic performance was also examined. Genetic similarity measures were poor predictors of the near-homozygous progeny performance and general and specific combining ability effects for three agronomic traits in the recurrent selection programme. In addition, there was no apparent trend for preferential selection of progeny from either more similar or more diverse parents within a given cycle of selection. The overall reduction in genetic diversity in this population has not affected selection response for grain yield. Variability remaining at important loci or de novo variation are possible explanations for the continued selection progress.     

Genetic-diversity assessed by microsatellites in tropical maize populations submitted to a high-intensity reciprocal recurrent selection

BR-105 and BR-106 are important tropical maize populations, which were submitted to a high-intensity reciprocal recurrent selection, generating the IG-3 and IG-4synthetics. Using 30-microsatellite loci,we measured and compared the genetic diversity of these populations and their synthetics. The populations did not differ significantly regarding the amount of genetic diversity. As a consequence of selection, genetic variability losses, in terms of mean number of alleles per locus,proportion of polymorphic loci, and gene diversity did occur and were greater in the synthetic IG-3 than IG-4. In the synthetics, the number of loci in adherence to Hardy-Weinberg proportions was superior to that observed in the populations. The Wright's mean fixation index was higher than the mean value expected for outcrossing species (5%) indicating as light excess of homozygotic individuals in both populations. The genetic distances confirmed the favourable effects of one cycle of recurrent selection, as the synthetics became more isolated in comparison to the original populations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Application of the concepts of test value and varietal value to the study of genetic advance in recurrent selection: A synthesis

Summary The concepts of varietal ability and value in test of a genotype or group of genotypes are specified according to the type of variety (clones, hybrids, lines, synthetics) and the testing system (phenotype, S₁, General combining ability (GCA), line value, General synthesizing ability (GSA)). Varietal ability and value in test are to be considered as any quantitative characters for which it is possible to define additive effects, dominance effects and epistasis. For diploids in recurrent selection, only additive effects and additive× additive epistasis for varietal value contribute to genetic advance. Genetic advance depends on the covariance between the testing value of the parents and the test value of the progeny by intercrossing. Such a parent-offspring covariance is, in the absence of epistasis and inbreeding, one half of the covariance between additive effects in test of the parents and additive effects in test of their progenies. This allows one to express directly the genetic advance in varietal value according to the type of variety and to the testing system used in recurrent selection. The case of biallelic populations is considered; it allows some discussions on the relative values of components of variance of varietal and test values.     

Reciprocal recurrent selection applied to Coffea canephora Pierre

Summary Genetic parameters were estimated in trials planted in connection with a reciprocal recurrent selection programme applied to Coffea canephora in Côte d'Ivoire. Narrow sense heritability values, estimated from a breakdown of the analysis of variance, were high for architectural characters (0.22 to 0.78) and medium for tree vigour (0.13 to 0.40) and for bean weight (0.15 to 0.28). Calculated heritability values for the first two harvests were high (h²>0.7), but much lower for the following harvests (less than 0.2). Heritability estimates by parent-offspring regressions gave variable results, though they were often similar to those obtained by variance analysis, especially for bean weight. Phenotypic and genetic correlations between tree vigour traits and productivity were high. Bean weight was not correlated with vigour or production. Canopy diameter of 4 year-old trees was closely correlated with cumulated productivity from 2 to 5 years in the absence of development competition between trees, but the correlation was low if trees were in competition with each other. The consequences of the results for the choice of characters to be selected among intergroup hybrids or the parents of both populations are discussed.     

Optimization of DH-line based recurrent selection procedures in maize under a restricted annual loss of genetic variance

Doubled haploid (DH) lines are increasingly being used in commercial hybrid maize (Zea mays L.) breeding. They allow for various quantitative genetic and logistic advantages provided that they are implemented in efficient and optimally allocated breeding procedures. In the present study, a new software was applied to optimize two recurrent selection (RS) schemes for hybrid maize breeding based on DH lines under a restricted annual budget and an upper limit for the relative annual loss of genetic variance. This software maximizes the expected gain from selection in general combining ability by means of quantitative genetic model calculations. Optimization results are compared for one, two, and three stages of testcross selection under different assumptions regarding the evaluation of lines per se and the annual budget. Results show that the optimum allocation of technical and budget resources to the individual steps of an RS program and the efficiency of alternative RS procedures are decisively determined by the number of selection stages. Under standard assumptions, one-stage selection was superior to two- and three-stage selection. Thus, shortening the length of an RS scheme considerably increases its efficiency. By intercrossing a reduced number of selected lines for starting a new RS cycle, the short-term response to selection may be increased, but the population size and, thus, the selection limits in the long run are diminished. Therefore, fair comparisons of alternative RS procedures require to define the intended time span for maximizing the genetic gain from RS and to restrict the relative annual loss of genetic variance accordingly.     

Continued response through seven cycles of recurrent selection for grain yield in oat (Avena sativa L.)

Long-term selection experiments provide germplasm to study the effects of selection in a closed population. Recurrent selection to enhance grain yield in oat has been ongoing at the University of Minnesota since 1968. The objectives of this study were: (i) estimate the GCA and SCA effects for three agronomic traits in the seventh cycle of selection, (ii) assess the effect of the current methods of selection on parental contribution and unselected traits, and (iii) determine the direct and indirect responses to seven cycles of recurrent selection for grain yield. Progeny of the Cycle 6 parents and parents for Cycles 0 through 7 were grown in two separate tests. Grain yield, heading date and plant height were evaluated in each test. Grain yield was increased by 21.7% after seven cycles of selection. Evaluation of Cycle 6 progeny showed that GCA effects were significant for all three traits studied, and SCA effects were significant only for grain yield. Four Cycle 6 parents did not have any progeny selected as Cycle 7 parents. Results from this study indicate that long term recurrent selection has continued to increase grain yield. Alternative selection strategies may be necessary to maintain the genetic variability in this population, particularly when improvement of secondary traits is required. This revised version was published online in August 2006 with corrections to the Cover Date.     

Methods of increasing short term response to full-sib family recurrent selection in small populations

Accelerated recurrent selection (ARS), in which selection is carried out on the predicted value of the progeny rather than on the observed performance of the parents, has been proposed as a method of increasing response to selection and of reducing cycle time. ARS schemes based on test cross evaluation of full-sib families have been compared by stochastic computer simulation. The difference in genetic and economic time scales is emphasised, with the economic long term (21 years) being only 21 or fewer cycles of selection. ARS schemes are shown frequently to offer improvements over standard recurrent selection methods under these circumstances, since they allow more cycles of selection in a given time frame. Schemes with very low effective population sizes often give the greatest response to selection over the time scales considered here. It is suggested that evaluation of cumulative responses to selection over defined periods of time, either by Monte Carlo simulation or by stochastic theory, is the best method of ranking alternative selection schemes. Evaluating response to selection by deterministic methods, or by attempting to take drift into account by evaluating schemes with identical effective population sizes can be misleading. This revised version was published online in August 2006 with corrections to the Cover Date.     

Divergent recurrent selection for cold tolerance in two improved maize populations

Maize (Zea mays L.) production has significantly expanded into very short-season environments where germination and growth in cooler environments is essentially a pre-requisite. Therefore, an important goal for maize breeders is to improve local germplasm sources of inbred lines that are able to grow under these challenging conditions. The objective of this research was to evaluate direct and correlated responses in two improved early maturing maize populations [NDSCD(M)C10 and BS22(R)C7] after two cycles of S₁ and full-sib intra-population recurrent selection for cold tolerance. The S₁ and full-sib progenies were obtained by self-pollinating 100 random plants and by intercrossing 200 random plants, respectively. Ten percent of the families were selected, based on an index that included emergence percentage, seedling vigor, and root lodging percentage, and recombined at the same time in a summer nursery based upon data across northern North Dakota locations. The essential benefit of this breeding methodology was to achieve one year per cycle of selection based upon progenies. However, direct response to selection was not significant while some correlated responses were significant. We decided to report these results in order to encourage other scientists the evaluation of additional sources of germplasm, the screening at various dates, and the selection of target environments with more intensive cold stress before initiating long-term selection programs for cold tolerance. In addition, further research on the current and alternative long-term selection methods for cold tolerance is recommended for continuous genetic improvement of advanced cycles in the northern U.S. Corn Belt. Part of the thesis submitted by B. Sezegen in partial fulfillment of the requirements for a MS degree at North Dakota State University.     

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.     

Five cycles of recurrent selection for increased essential oil content in East Indian lemongrass: response to selection, and effects on heritabilities of traits and intertrait correlations

Five cycles of phenotypic recurrent selection for increased essential oil content were carried out in East Indian lemongrass, Cymbopogon flexuosus. In each cycle, the top 5% of plants for essential oil content were selected and their ramets were planted in isolated polycross blocks to produce the seed of the next cycle. Response to selection for essential oil content and its effect on three unselected traits, leaf yield, dry matter content in leaves and citral content in the essential oil, were determined by evaluating C₀‐C₅ populations in a replicated trial. Genetic variation, heritability estimates and intertrait correlations for essential oil content, leaf yield, leaf width, tiller number and citral content in the essential oil in C₄ were determined, by evaluating 40 clones and their half‐sib progenies produced from 40 randomly selected C₄ plants, in a replicated experiment. These were compared with those determined in C₀ earlier. The mean essential oil content increased from 0.66% in C₀ to 1.67% in C₅ (i.e. by about 31% per cycle over C₀). Selection for essential oil content did not affect leaf yield and its effect on dry matter content in leaves was marginal. The first three cycles of selection for essential oil content did not affect citral content in the oil but two further cycles significantly decreased citral content. Heritabilities and intertrait correlations between all traits studied, except citral content and essential oil content, were similar to those found in C₀.     

Use of index selection in recurrent selection programs in maize

Summary Phenotypic and genotypic correlations were examined for four traits in seven populations of maize (Zea mays L.) undergoing recurrent selection. Correlations among grain yield and percentage of grain moisture, root lodging, and stalk lodging were low (|r|<0.3) except for the correlation between grain yield and stalk lodging, which was high and negative. The phenotypic and genotypic correlations agreed well from cycle to cycle within populations. Variation of correlations among populations was not significantly larger than variation among cycles. Heritabilities of these traits generally were high (h²>0.5). Two indices, one that used heritabilities as index weights and one that used relative economic weights (base index) as index weights, were compared with the Smith-Hazel index (optimim index). Relative efficiencies of the two indices, in terms of predicted gains for the individual traits and the composite trait, compared with the Smith-Hazel index, were high. The use of an index in which heritabilities were used as index weights was recommended because:1) the heritabilities were the same as the optimum weights when the traits were uncorrelated, and for the data examined the correlations were low; and 2) heritabilities were computed in routine data analyses and were available at no additional cost.Joint contribution: USDA-SEA-AR and Journal Paper No. 10152 of the Iowa Agriculture and Home Economics Exp. Stn., Ames, Iowa. Project 2194.     

Reciprocal recurrent selection applied to Coffea canephora Pierre. III. Genetic gains and results of first cycle intergroup crosses

Breeding of the allogamous Coffea canephora species in Côte d'Ivoire is based, from 1984 onward, on a reciprocal recurrent selection scheme exploiting the good performance of crosses between two genetically distinct populations (‘Guinean’ and ‘Congolese’). Results are presented here on intergroup cross progenies evaluated on individual plant basis over a six year period in three fully randomized trials planted in 1985 and 1986. Yields of the bi-parental crosses varied from 16 to 1407% of the mean of the two commercial clonal cultivars and the most productive crosses were also the most vigourous. Selection of individual trees and cross progenies, based on a combined individual-family index, was carried out for high yield, good vigour of young plants and moderate canopy diameter of adult trees. Expected genetic gains, as estimated in relation to the mean of all crosses, for selection of 5% of the best trees, were high for yield (over 60%), moderate for young plant vigour (14 to 18%), and, as intended, low for canopy diameter. In addition, selection by independent culling for leaf rust resistance and bean weight was carried out. As a result, 6 promising crosses with yield similar or higher than those of the control cultivars, were selected as candidates for commercial use, as well as 65 individual trees, within 19 different crosses. Expected genetic gain for yield for these selected trees is about 60% compared to the average yield of the most productive control clone. Continuation of the programme is discussed, with specific attention to further progress expected for quality traits.     

Analysis of sub-populations of rapid-cycling Brassica rapa following recurrent bi-directional selection for cotyledon size

Recurrent selection programmes use one or just a few selection criteria, however other indirect traits may be unpredictably changed in a population. This study was conducted to determine the indirect effects of 10 cycles of bi‐directional recurrent selection for cotyledon size in the model system of rapid‐cycling Brassica rapa. Eight sub‐populations (four large‐ and four small‐cotyledon sizes) were phenotypically and cytologically evaluated. Each sub‐population was measured by five phenotypic traits. Ploidy and pollen viability were studied and compared with the initial population. Total weight was significantly different in broad and bottleneck‐1 sub‐populations. Total cell number exhibited statistically significant differences in broad and bottleneck‐1 sub‐populations, while cell number per unit area exhibited statistically significant differences in broad, bottleneck‐2 and bottleneck‐3 sub‐populations. Decreases in pollen viability in comparison with the base population were observed in three sub‐populations. Among the eight sub‐populations studied, the most significant phenotypic differences were observed within broad sub‐populations. Based on the above, it is possible that bi‐directional recurrent selection for cotyledon size may have been a result of indirect selection for two processes, endoreduplication and cell division.     

Sporophytic and gametophytic recurrent selection for improvement of partial resistance to Alternaria leaf blight in sunflower (Helianthus annuus L.)

Alternaria leaf blight, caused by Alternaria helianthi Hanf., is one of the most important diseases of sunflower causing significant yield losses in several tropical countries. Yet, so far, only partial resistance for the disease has been discovered in the germplasm through conventional sporophytic selection. Therefore, the main objective of the present investigation was to compare sporophytic and gametophytic recurrent selection with the aim to enhance the level of resistance to Alternaria leaf blight. The base population was synthesized by random mating three populations- two interspecific derivatives involving different species of Helianthus and one germplasm accession based on their partial resistance to disease incidence. The base population was subjected to 1-2 cycles of both sporophytic and gametophytic selection. The gametophytic selection was practiced by applying pathogen culture filtrate to the stigma and style one hour before pollination. The selection response was measured by scoring the percent disease index at flowering, 15 days after flowering, and at physiological maturity and by quantifying economic yield gain. A significant reduction in mean per cent disease index values and a gain in seed yield were observed for both the types of selection cycles, but more so for gametophytic selection. The populations improved through gametophytic selection appear to be more promising as the pollen selection allowed the selection of rare favorable allelic combinations that would hardly be detected at the sporophytic level. A combination of gametophytic selection and conventional sporophytic selection should be considered as an effective tool in population improvement programs to achieve higher levels of resistance in relatively short time.     

Changes in morphological and physiological traits associated with recurrent selection for grain yield in maize

Summary Experiments were conducted during the growing seasons of 1975 and 1976 to determine changes in morphological and physiological traits associated with recurrent selection for grain yield in maize (Zea mays L.). Four variety hybrids, BSSS(R)CO x BSCB1(R)CO, BSSS(R)C7 x BSCB1(R)C7 [from a reciprocal recurrent selection program involving Iowa Stiff Stalk Synthetic (BSSS) and Iowa Corn Borer Synthetic #1 (BSCB1)], BS12CO x B14A, and BS12C6 x B14A [from a half-sib selection program involving the open-pollinated variety Alph (BS12) and the inbred tester B14A] were grown at 59 300 and 98 800 plants/ha near Ames, Iowa We obtained data on CO₂-exchange rate (an estimate of photosynthetic rate), grain yield, grain-yield components, flowering dates, maturity traits, light interception and use, shelling percentage, harvest index, and various other plant traits.CO₂-exchange rate did not change appreciably with recurrent selection for yield. Grain yield per hectare and per plant were larger for the improved than unimproved hybrids. Grain-yield components did not change significantly with recurrent selection. Kernel weight of BSSS(R)C7 x BSCB1 (R)C7, however, was larger than that of BSSS(R)C0 x BSCB1(R)C0. Pollen-shed-to-silking interval was shorter for the improved than the unimproved hybrids, and grain-filling duration was longer in C7 x C7 than in C₀ x C₀ of the reciprocal recurrent selection program. Furthermore, improved hybrids were characterized by smaller tassels and more upright canopies. Usually, plant traits and leaf-area-related traits were similar for all hybiids.Although dry-matter productivity was similar for all hybrids, those that were improved by recurrent selection produced more grain per unit leaf area and per unit light interception. Also, BS12C6 x B14A was characterized by a higher harvest index than BS12CO x B14A.We concluded that the source (i.e., photosynthetic capacity) was not limiting grain yield in BSSS(R) x BSCB1(R) and BS12. Increased grain yields that resulted from recurrent selection were consequences of longer grain-filling duration for BSSS(R) x BSCB1(R) and increased translocation of photosynthate from source to sink for both BSSS(R) x BSCB1(R) and BS12.Journal Paper J-8953 of the Iowa Agriculture and Home Economics Exp. Stn., Ames, Iowa. Project No. 2152.     

Biometrical analysis of alternative plot types for selection in rye

Summary Maximizing the efficiency of selection for yield and baking quality of winter rye (Secale cereale L.) requires reliable knowledge on the pertinent population parameters. This study reports estimates of variances, heritabilities and genetic correlations from a) large (5 m²) drilled (LD) plots, b) micro drilled (MD) plots, and c) one-row plots of 4 spaced plants (SP). Thirty eight single crosses of rye were grown at two locations for two years in LD, MD and SP plots replicated 2, 4, and 6 times, respectively. Genotypic differences were significant in all plot types for all agronomic and quality traits, although estimates of genetic variances were smaller in LD than MD or SP plots for grain yield, 32-spike weight, and kernels per spike. Heritabilities on an entry mean basis (0.62 to 0.95) were similar among plot types, but on a single plot basis estimates were lower for SP than for MD or LD plots. Genetic correlations were high between all three plot types, with correlations between LD and SP plots being approximately 1.0 for falling number, thousand-kernel weight, and bloom date, 0.90 for grain yield, kernels per spike, test weight and height, and 0.70 for tiller number. Based on the high heritabilities achieved in small plots and the close agreement between MD- or SP- with LD-plot performance we conclude that greater use should be made of small plot types when selecting for yield and quality in winter rye.