The efficiency of between and within full-sib family selection in a recurrent selection programme in sugar beet (Beta vulgaris L.)

The efficiency of between and within family selection in a full-sib family recurrent selection programme was investigated using data from three cycles of recurrent selection. Correlations between mid-parent values and offspring were high for sugar content and juice purity characters, but low for root weight and sugar yield. This suggests that single root selection within full-sib families is effective in the improvement of sugar content and juice purity, but ineffective for root weight and sugar yield. Root weight and sugar yield will respond better to between full-sib family selection. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Divergent full-sib recurrent selection for germination at low temperature in a maize population

Summary Four cycles of divergent full-sib recurrent selection for the ability to germinate at low temperature were conducted in a maize (Zea mays L.) F₂ population. The selection criterion was the high (H) or the low (L) value in algebraic terms of the difference (DG) between germination percentage at 9.5°C (G9.5) detected 19 days after sowing and germination percentage at 25°C (G 25) seven days after sowing; both traits were evaluated in a controlled environment (germinator). Direct and correlated responses estimated during the course of selection were in accordance with those evaluated at the end. Selection for H led to populations with higher DG values, while the reverse was noted for L; differences between H and L populations increased in successive selection cycles, though divergence tended to level off. Selection for H also resulted in higher G 9.5 (day 19), shorter germination time and more flinty kernels, while selection for L led to responses in the opposite direction as well as to a lower G 9.5 detected 37 days after sowing (i.e. at the end of germination). In contrast, responses were negligible for G 25 and varied erratically from one cycle to another for kernel weight.Abbreviations DG (G 9.5–G 25) - FS full sib - GI germination index - G 9.5 germination percentage at 9.5°C - G 25 germination percentage at 25°C - H high DG value in algebraic terms - KT kernel type - KW kernel weight - L low DG value in algebraic terms     

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.     

An extension of the Smith model for quantitative genetic analysis of selection response under recurrent selection

Selection and random genetic drift are the main forces affecting selection response in recurrent selection (RS) programmes. The correct assessment of both forces allows a better comparison of the efficiency of different RS schemes. The objective of this study was to extend the population diallel analysis proposed by Hammond and Gardner in 1974 and the model proposed by Smith in 1979 with full consideration of inbreeding depression due to random genetic drift. The effect of random genetic drift is expected to be large, particularly in studies with many selection cycles and/or high rates of inbreeding. Therefore, the extension of the population diallel allows a better assessment of the selection response in RS programmes.     

Effects of recurrent selection for grain yield on plant and ear traits of five maize populations

Summary Maize (Zea mays L.) breeders are interested in the effects of recurrent selection for grain yield on other traits. Changes in plant traits could alter agronomic acceptability of the populations under selection, and observed improvements in grain yield could be explained by changes in ear traits. We evaluated changes in combining ability for plant and ear traits of BS10(FR), BS11(FR), BSSS(R), BSCB1(R), and Lancaster Surecrop associated with recurrent selection for grain yield.Recurrent selection procedures generally did not change plant and ear heights or date of silking of testcrosses of populations or of the population crosses, BS10(FR)×BSS11(FR) and BSSS(R)×BSCB1(R). Grain yield improvements, however, were associated with increases in ear-sink size.Journal Paper No. J 9517 of the Iowa Agriculture and Home Economics Exp. Stn., Ames, Iowa. Project 2152.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Feasibility of breeding male-sterile populations for use in developing interpopulation hybrids of pearl millet

Inter‐population hybrids of pearl millet, Pennisetum glaucum (L.) R. Br., have a substantial grain yield advantage over open‐pollinated varieties that makes them an appropriate and economically viable proposition for many African agricultural situations, provided that stable male‐sterile populations can be developed for use as seed parents. The objective of this research was to examine the feasibility of breeding stable male‐sterile populations, using the d₂ dwarf version of Nigerian Composite NCD₂ and the A₄ cytoplasmic‐nuclear male sterility system as a test case. Results showed that two cycles of recurrent selection for sterility maintenance ability led to the development of a fully effective maintainer version of NCD₂. There was no significant difference between the original C₀ cycle bulk and the C₃ cycle bulk (developed from the third and final cycle of recurrent selection) for grain yield and other agronomic traits. The male‐sterile population at the third backcross stage, developed from the maintainer version of NCD₂, had as high a level of stable male sterility as the A¹ system commercial inbred male‐sterile line 841A₁. Thus, it is concluded that with the use of the A₄ cytoplasmic male‐sterile system, it would be possible rapidly to develop a maintainer version of any population without detrimental effects on grain yield and agronomic traits. Male sterility of populations developed from these maintainers will be highly stable, paving the way for their effective utilization as seed parents in breeding inter‐population hybrids.     

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.     

Phenotypic recurrent selection in ideotype breeding of pinto beans

Summary Plant growth habit (type) plays a major determining role in the acceptance of a dry bean cultivar by commercial growers. Under the threat of wet fall conditions in the midwestern U.S., growers show a preference for upright plant types-I and II over the more prostrate type-III growth habit of commercial pinto cultivars. The prostrate habit creates considerably more risk of harvest losses and potential of white mold (Sclerotinia sclerotiorum) development during a wet growing season.The breeding strategy being utilized to develop better adapted pinto bean cultivars for the humid midwest is based on the concept of ideotype breeding which has been successfully applied to small seeded navy and black beans. A system of phenotypic recurrent selection using S₁ selection is being followed in order to recombine desirable traits from two diverse germplasm pools. One germplasm pool possesses the desirable architectural plant form and disease resistance traits of the small-seeded types while the medium-seeded germplasm source possesses the desirable seed size, shape and color characteristics of the pinto class. Since a repulsion phase linkage appears to exist between type-II architecture (architype) and medium seed size (40 g/100 seeds), recurrent selection is shown to be the most effective procedure for breaking up undesirable linkage groups and for increasing the frequency of desirable genes, thus enhancing the chance for the desirable genetic recombinations to occur.Journal Article No. 11667 of the Michigan Agricultural Experiment Station.     

Computer simulation of a selection strategy to accommodate genotype environment interactions in a wheat recurrent selection programme

Multi-environment trials (METs) are used in plant breeding programmes to evaluate genotypes (lines/families) as a basis for selection on expected performance (yield and/or quality) in a target population of environments (TPE). When a large component of the genotype environment (G × E) interactions results from crossover interactions, samples of environments in METs that deviate from the TPE provide a suboptimal basis for selection of genotypes on performance expected in the TPE. To adjust for the negative effects of these deviations, a selection strategy that weights the data from the MET according to their expected frequency of occurrence in the TPE (i.e. a weighted selection strategy) was investigated. Computer simulation methodology was used to obtain preliminary information on the weighted selection strategy and compare it to the traditional unweighted selection strategy for a range of MET scenarios and G × E interaction models. The evaluation of the weighted selection strategy was conducted in context with the germplasm enhancement programme (GEP) of the Northern Wheat Improvement Programme in Australia. The results indicated that when the environments sampled in the MET matched those expected in the TPE, the unweighted and weighted selection strategies achieved a similar response to selection in the TPE. However, when the environments sampled in the MET did not match the expectations in the TPE and a large component of the G × E interactions resulted from crossover interactions, the weighted selection strategy achieved a greater response to selection in the TPE. The advantage of the weighted strategy increased as the amount of crossover G × E interaction increased or fewer environments were sampled in the METs.     

Inheritance in seed size of alfalfa: Quantitative analysis and response to selection

Seed size in alfalfa (Medicago saliva L.) has been positively correlated with seedling vigour and early growth, but there were few published reports on inheritance or selection for this trait. The objective of this research was to estimate components of genetic variance for inheritance of alfalfa seed size and determine the most efficient selection method. Components of genetic variance were estimated on seed and pollen plants of ‘BIC-7-WH’ and their progeny arranged in a North Carolina Design II mating design under controlled environmental conditions. Three selection methods, differing in parental control and selection pressure, were used to determine selection response. The seed parent genotype had a major role in determining alfalfa seed size, but the genotype of the seed had no influence. For genetic studies, pollen and seed parent effects on seed size should be measured on seed harvested from progeny plants. Seed size was controlled by additive and non-additive components of genetic variance. Heritability for seed size was 41.3%. Selection for seed size was effective and a significant shift for larger and smaller seed was attained after one cycle of selection.