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.     

Inheritance of groat-oil content and high-oil selection in oats (Avena sativa L.)

Summary The potential for breeding for high groat-oil content in oats was investigated by (a) conducting generation means analyses on data from three matings among adapted Avena sativa L. cultivars, (b) practicing one cycle of phenotypic recurrent selection in a segregating population derived from eight species backcrosses (Avena sativa x (A. sativa x A. sterilis)) among 24 parents, and (c) identifying transgressive segregates from interspecific (A. sativa x A. sterilis) matings.Additive gene action was the most important component in explaining the variation among generation means for groat-oil content. Dominance and epistatic interactions involving dominance were not significant in any mating. Significant residual genetic variation occurred in one mating, even after additive, dominance, and three digenic interactions were fitted. The importance of additive genes action implies that desired allelic combinations for high groat-oil content can be obtained in pure-line cultivars.One cycle of phenotypic recurrent selection using single plants as the selection units resulted in a genetic gain of 1.7 to 2.1% in groat-oil content. Individual plants selected for initiating the second cycle had from 9.5 to 12.6% groat oil.Over all 12 interspecific matings, the F₂ progeny means were similar to the midparent values. Only two were significantly deviant. Transgressive segregates for high and low groat-oil content from these matings provided evidence that A. sterilis possesses alleles for high and low groat-oil content that are different from those in the gene pool of cultivated oats.Journal Paper No. J-11340 of the Iowa Agric. and Home ECon. Exp. Stn., Ames, Iowa 50011. Project 2447. This study was supported in parts by grants from the Iowa Committee for Agricultural Development and the International Harvester Company.     

Morphology, Cytology and Fertility of Avena sativa L. ×A. magna Murph. et Terrell Amphiploids

The genotypes of Avena sativa were crossed to A. magna Murph. et Terrell, from the sterile pentapliod F₁ amphiploids were successfully developed after colchicine treatment. The C₁ plants were observed for their morphology, cytology and fertility. These plants were characterized by longer and broader glumes, swollen pedicel attachment, pubescent lemmas, bold seeds and persistent spikelets. Other characters, viz., final plant height, rachis length, peduncle length and spikelets per panicle were intermediate. All plants hail the chromosome number 2n = 70 or close to. 7C. Due to stickiness of chromosomes, meiosis of only two decaploid plants could be studied in detail. The average chromosome Association was 1.3 I + 28.56 II + 1.4 III + 1.77 IV + 0.07 VI and 0.13 I + 28.7 II + 0.14 III + 2.09 IV + 0.59 VI; but in some cells up to 35 bivalents were observed. Seed set as well as pollen fertility and size exhibited great variation.     

Chromosomal Stability in the Backcross Progenies of Pentaploid Hybrids between Avena sativa L. and A. maroccana Gdgr.

In a backcrossing programme to transfer desirable characters from wild Avena maroccana Gdgr. to cultivated oats, A. sativa L., meiotically stable plants in BC₁F₃ and BC₂F₂ progenies were isolated. The recovery of stable genotypes with 2n = 6×= 42 chromosomes indicated that two backcrosses are enough for such a programme. The cytological observations in various backcross generations are presented and discussed.     

Association of gene for grain colour with monosome XIV of Avena sativa L.

Summary The F₂'s from crosses between Dyfed (S.240; a black grained cultivar) and monosomics of Sun II (non-black cultivar) were analysed. With the exception of F₂ progenies involving monosomes, II, VI, VII, X, XI, XII, XIII and XIV, all other deviated from the genetic ratio 3 black: 1 non-black. The cytological examination of the non-black individuals belonging to those families that did not deviate from the 3:1 ratio showed that they were either 41- or 42-chromosome plants. However, non-black individuals involving monosome XIV were 40-chromosome plants. The gene conferring the black colour to the grains in Avena sativa cv. Dyfed (S.240) is located on chromosome XIV.Department of Plant Breeding and Genetics, Sind Agriculture University. Tando Jam. Pakistan.     

Chromosomal location of genes for resistance to powdery mildew in cultivated oat (Avena sativa L.). 1. Gene Eg-3 in the cultivar 'Mostyn'

The cultivar 'Mostyn' of hexaploid oat ( Avena sativa L.) possesses a dominant gene conferring resistance to powdery mildew caused by Erysiphe graminis avenae. The resistance gene, designated Eg -3, was derived from the wild oat Avena sterilis L. var. ludoviciana. Monosomic analysis revealed that this gene was located on chromosome 17, which corresponds to the missing chromosome in the 'Kanota' monosomic line Kl1. This chromosome is associated with the homoeologous group 1 of the Triticeae.     

Correlated responses of fatty acid composition, grain quality, and agronomic traits to nine cycles of recurrent selection for increased oil content in oat

Increases in the groat-oil content of oat (Avena sativa L.) increase the energy value of the grain and improve the feasibility of extracting oat oil for use as a vegetable oil. Nine cycles of recurrent selection for greater groat-oil content conducted in a genetically broad-based oat population resulted in dramatic increases in groat-oil content. Our objectives were todetermine if selection for greater groat-oil content affected fatty acid composition, grain quality traits (test weight and seed weight), or agronomic traits (straw yield, biomass, harvest index, heading date, and height). We evaluated 100 random lines from the base (C0) population and each of the nine selection cycle populations in three environments in order to estimate means, genetic variances, heritabilities, and genotypic and phenotypic correlations of grain quality and agronomic traits. We also evaluated 20 random lines from each population to estimate changes in fatty acid contents. Oleate and stearate contents increased over cycles of selection, as did the ratio of unsaturated to saturated fatty acids. Palmitate, linoleate, and linolenate contents and all grain quality and agronomic traits except harvest index decreased over cycles of selection. There was no evidence for reduced genetic variance or heritability in C9 for any trait, but the genotypic and phenotypic correlations between agronomic traits and oil content fluctuated over cycles. Selection for increased groat-oil content improved oil quality but reduced grain quality and agronomic performance of the population. This revised version was published online in August 2006 with corrections to the Cover Date.     

The Performance of Pea (Pisum sativum L.) and its Role in Determining Yield Advantages in Mixed Stands of Pea and Oat (Avena sativa L.)

On a brown warp soil (Fluventic Eutrochrept) near Goettingen, Germany, conventional leafed pea ( Pisum sativum L. cvs Messire and Bohatyr) and semileafless types (cvs Profi, Juno and Azur) were grown in mixed stands together with oat ( Avena sativa cvs Alf and Lutz) in substitutively designed experiments from 1995 to 1997. Oat was the dominant component. Crowding coefficients for oat averaged 7.4. No relationship could be detected between the crowding coefficient of oat and any yield advantage from the mixture. Crowding coefficients for pea varied substantially, between 0.1002 (Juno and Alf in 1996) and 0.2979 (Bohatyr and Alf in 1996). Crowding coefficients for semileafless pea cultivars were smaller than for conventional leafed types. The yield advantage of the mixture increased as the crowding coefficient of pea increased. The maximum yield increase for the mixture was achieved when the relative yield total (RYT)=1.17 or + 11 dt grain DM ha^–1 for mixtures of the long-strawed conventional leafed cultivars Bohatyr and Alf (in 1996). The crowding coefficients of pea were positively correlated with the level of symbiotically fixed N₂ in the mixed stands. When N₂ fixation with mixed cropping was about 30 kg N ha^–1, RYT was unity. Increasing symbiotic N₂ in the mixtures resulted in increasing yield advantages in the mixture. Short-strawed pea cultivars seem unsuitable for mixing with oat. Plant height of pea appeared to be more important than plant leaf type. Accordingly, mixtures containing the long-strawed semileafless pea cultivars Profi and Alf were more successful. It is concluded that increased competitiveness of the pea component in the mixture with oat entails increasing the level of symbiotic N₂ fixation including resource complementarity and thus yield advantage in the mixed stands.     

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.     

Phenotypic recurrent selection in common bean (Phaseolus vulgaris L.) based on performance of S2 progenies

Summary The effect of a recurrent selection procedure for improving seed yield per plant was evaluated in common bean. Progress was registered after three cycles of recurrent selection with S2 progeny evaluation performed on the grain yield/plant. Gain from selection was estimated by field testing 45 random S2 lines from each of the C0, C1 and C2 populations. Response to selection for seed yield/plant was 6 g from C0 to C1 and 4.4 g from C1 to C2. The seed yield improvement was due principally to an increase in pods/plant; little contribution was provided by the 1000-seed weight and seeds/pod traits. Broad-sense heritability and genotypic variance for grain yield remained high in all cycles which suggests further gain from additional selection cycles. Grain yield/plant is significantly correlated to pods/plant and seeds/pod traits in all cycles, indicating that plants with favourable combinations of these traits could be identified. The proposed recurrent selection procedure appears to be effective in improving the population and extracting superior genotypes for varietal development.     

Identification of New Genes for Mildew Resistance of Barley at the Mia Locus in Lines Derived from Hordeum spontaneum

Genetical studies have been conducted with lines derived from H. spontaneum showing that these lines carry dominant mildew resistance genes located at or near the Mla, locus. The resistance spectra of the lines ‘RS170—10 × Piccolo A’, ‘Diamant × 1B-20’, ‘RS1—8 × Piccolo E’ and ‘Diamant × 1B-151’ obtained from 10 European and Israeli isolates differ from previously-identified Mia alleles. Therefore, it is suggested that these genes should be designated as Mla 25, Mla 26, Mla.27 and Mla 28, respectively. In addition, the RFLP-patterns of these lines and their crossing parents were studied by hybridization with probes MWG 1H036, MWG 1H060 and MWG 1H068, which are very closely linked to the Mia locus. Two double crossover events have been identified. The use of RFLP markers for the identification of mildew resistance genes is discussed.     

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.     

Comparison of evaluation methods for selection of resistance to Fusarium head blight in a recurrent selection programme in wheat (Triticum aestivum L.)

The objectives of this study were to compare efficiency of evaluation for resistance to Fusarium head blight (FHB) under two inoculation methods in a recurrent selection programme. Fifty selected homozygous F₅ fertile lines, from each of five cycles (C0, C1, C2, C3 and C4) of recurrent selection, and two control cultivars were evaluated in a split-plot design in 1995 and 1996 under the soil-surface inoculation with Fusarium graminearum-colonized kernels and the single-floret inoculation with ascospore suspension. Comparison of the two inoculation methods using means, ranges, coefficients of variation, heritabilities and correlations among infected spikelet rate (ISR), reaction index (RI) and disease index (DI) indicated that FHB resistance could be evaluated with similar accuracy and precision using either of the two inoculation methods. Regressions of disease scores in the soil-surface inoculation on disease scores in the single-surface inoculation were positive and highly significant, showing a strong relationship between both inoculation methods for FHB resistance. The percentage of lines with similar performance for FHB disease scores in both inoculation methods was high. The soil-surface inoculation and single-floret inoculation appear to be useful techniques for evaluating numerous individuals of segregating population and screening advanced homozygous lines for FHB resistance in a recurrent selection programme in wheat, respectively.     

Predicting the relative effectiveness of direct versus indirect selection for oat yield in three types of stress environments

Summary In breeding crop varieties for stress environments, it must be decided whether to select directly, in the presence of stress, or indirectly, in a nonstress environment. The relative effectiveness of these two strategies depends upon the genetic correlation (r _g ) between yield in stress and nonstress environments and upon heritability in each. These parameters were estimated for grain yield of 116 random oat lines grown in nonstress, P-deficient, N-deficient, and late-planted environments. Estimates of r _g between yield in nonstress and yield in P-deficient, N-deficient, and late-planted environments were 0.52±0.24, 1.08±0.16, and 0.06±0.24, respectively. No consistent relationship between heritability and environment mean yield was observed. Direct selection in the presence of stress was predicted to be superior for yield in low-P and late-planted environments, but indirect selection in high-N environments was predicted to be as effective as direct selection in producing yield gain in low-N environments. These results confirm that neither high-yield environments nor environments in which the heritability of yield is maximized are necessarily optimum when the goal is to maximize yield gain in stress environments.Dep. of Agronomy, Iowa State Univ., Ames, IA 50011; USA, Journal Paper no. 13101. Project 2447.     

Bulked AFLP analysis for the assessment of genetic diversity in white clover (Trifolium repens L.)

The use of bulked leaf samples from individual plants for amplified fragment length polymorphism (AFLP) analysis was evaluated as a tool for assessment of genetic diversity in white clover (Trifolium repens L.). Bulking of leaf samples produced slightly simpler AFLP profiles compared to the combined profiles of individual plants from the same cultivar. Approximately 90% of bands which were present in individual plants were present in bulked samples of the same cultivar. The majority of those absent were rare bands, shared by less than 25% of individual plants. Replicate bulk samples gave almost identical banding patterns, demonstrating the robustness of the bulked AFLP technique. Cluster analysis of AFLP data derived from individual plants resulted in a phenogram similar to that produced from data derived from bulked samples of the same plants. AFLP analysis of bulked samples detected significant amounts of genetic variability among 52 cultivars and accessions with genetic similarity values ranging from 0.42 to 0.92. However, cluster analysis of AFLP data only partially reflected the geographic origin of cultivars and accessions and was not congruent with cluster analysis based on variation for morphophysiological characters. Bulked AFLP analysis provides a powerful tool for rapid assessment of genetic variability in white clover and may also be used for cultivar identification. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effect of moving mean covariance adjustments on error and genetic variance estimates and selection of superior lines in peanuts (Arachis hypogaeal L.)

Summary Effects of moving mean covariance adjustments on error control, genetic variance component estimates and selection were examined in nine trails in peanut breeding nurseries. Neighbourhood sizes of two to fourteen adjacent plots were used in two types of moving mean covariates: one based on adjacent plot yields and a second based on adjacent plot residuals after removal of replicate and treatment effects. Both covariate types reduced error variance, in one trial by up to 57%, although the covariate type giving the most error reduction varied among trials. Generally most error variance reduction occurred when six to eight adjacent plots were used.In some trials genetic variance component estimates following covariance adjustment were higher than those from unadjusted analyses and in other trials they were lower. As the neighbourhood size included in the moving mean increased from two to fourteen plots the genetic variance component estimate changed considerably in some trials. However, large rank changes did not appear to be associated with these genetic variance component changes as neighbourhood size had little influence on the selection of progenies. Also, genetic effects included in the yield covariate had little influence on selection of elite lines as essentially the same lines were chosen or rejected when residual covariates lacking these genetic effects wre used.     

Comparison of some selection procedures and objectives in faba beans (Vicia faba L.). I. direct and index selection for seed yield

Summary Starting with two heterogeneous F₂ populations, each based on 20 cross combinations, different selection procedures were applied for four consecutive selection cycles. Three of the selection procedures involved seed yield per plant (SYP): (i) direct selection for SYP (DSY); (ii) indexselection for SYP (ISY) with plant length, the ratio of the vegetative top to plant length, end of grain filling and duration of the generative stage as auxiliary characters and finally; (iii) independent culling levels selection (ICL) for the index described above and crude protein content. For ICL-selection the attention is focussed on SYP in this paper.For all three selection procedures, predicted and realized responses were calculated and expressed as percentage of a corresponding non-selected population. The intensities of selection differed over selection methods and generations, mostly due to practical constraints.As far as predicted responses are concerned, ISY-selection seemed to be slightly superior to DSY-selection and DSY-selection to ICL-selection. This order may partly be due to differences in selection intensity. However, none of the predictions was very accurate, but the predictions of ICL-selection seemed to be the most accurate. This has been ascribed also to the relative mild selection intensity applied in ICL-selection. It is concluded that a decision in favour of a complicated selection procedure like index selection based on predicted responses only cannot be defended. As far as realized responses are concerned, DSY-selection seems to be slightly superior to the other two selection procedures, as it is the only procedure that resulted in a more or less stable positive response to selection.The auxiliary characters, especially plant length, showed a marked correlated response to all three selection procedures. In case of direct selection for SYP this seemed paradoxical with the inefficiency of index selection, but, it may reflect the differences in the genuine genotypic correlations and the estimated genotypic correlations.     

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.     

Inter-and intraracial hybridization and selection for seed yield in early generations of common bean,Phaseolus vulgaris L.

Summary Five parents from each of four race groups of common bean (Phaseolus vulgaris L.) were hybridized to produce five crosses within each group. Also, five crosses were made for each of the six possible combinations among four groups. Parents, F₁ and F₂, and parents, F₂ and F₃ were evaluated for seed yield in 1990 and 1991, respecitively, at two locations in Colombia.Yield of parents belonging to Middle American races and crosses among them was higher than that of races of Andean origin. Positive correlations were found among the mid-parent value, F₁, F₂ and F₃. Also, the mid-parent value predicted the mean seed yield of all possible lines that could be derived from the F-generation in 42 out of 47 crosses. Four crosses, all between common bean races of Andean and Middle American origin, indicated a possible loss of favorable epistatic parental alleles. On average, mean yield of interracial F₁ hybrids was higher than that of intraracial ones.Positive heteroris (26.4%–123.8%) over the mid-parent in 31 crosses, and F₁s yielding higher (23.7%–91.8%) than the high parent in 20 crosses and yielding higher (22.1%–53.2%) than the highest control among all parents (MAM 13) in 12 crosses, were found. Heritability, estimated by the parent-offspring regression, ranged from 0.42± 0.07 to 0.49±0.04. Expected and realized gains from selection (at 20% selection pressure) ranged from 10.3% to 21.0% over the mean of F₁ hybrids and F₂ and F₃ population bulks.