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 parental lines of Indica hybrid rice (Oryza sativa L.) in China based on coefficient of parentage

Genetic diversity constitutes the raw material for plant improvement, and provides protection against genetic vulnerability to biotic and abiotic stresses. Diversity of parental lines of indica hybrid rice in China is not well‐characterized. The major objective of this study was to quantify genetic diversity of Chinese parental lines of hybrid rice via coefficient of parentage (COP). All 100 parental lines of hybrid rice widely used in hybrid breeding and commercial production during 1976–2003 were studied by COP analysis. The mean COP for the 100 parental lines was low (0.056), indicating a potentially high degree of diversity in Chinese hybrid rice breeding. Forty‐nine percent of all pairs of parental lines were completely unrelated by pedigree data. The low mean COP for the parental lines was attributed to a continual incorporation of exotic germplasm (wild rice, japonica and javanica etc.) into the genetic base over time, to the introduction of foreign germplasm from the Philippines (International Rice Research Institute), Korea, the United States, Thailand, and Guyana as breeding stock. The mean COP from 1976 to 1990 was twice as much as that from 1990 to 2003. Cluster analysis was an effective method to discriminate diversity, ten clusters were identified, and maintainer lines, restorer lines and other parental lines with special genetic background were clearly grouped. In addition, restorer lines were further divided into 11 sub‐clusters, which basically was in agreement with hybrid rice breeding. Among ten provinces, Hunan, Sichuan and Fujian were outstanding for breeding 54 of 100 parental lines in hybrid rice production, and the genetic diversity of parental lines in Fujian, Sichuan,Guangxi, Hunan and Jiangsu were all narrower than that in Hubei, Guangdong, Zhejiang and Jiangxi. The result of coefficient of parentage analysis for 100 parental lines may promote the management of parental diversity and hybrid rice breeding in China.     

Pedigree, RAPD and simplified AFLP-based assessment of genetic relationships among Avena sativa L. cultivars

Two molecular marker techniques: RAPD and simplified ^PstIAFLP have been compared in order to decide on, which technique is better suited to genetic characterization of oat (Avena sativa L.) cultivars. It was investigated, if the same pattern of variability is revealed by two approaches and whether the observed molecular variability reflects pedigree-based relationships. Polymorphic RAPD and ^PstIAFLP markers were sufficient to distinguish all analysed cultivars, demonstrating the usefulness of both methods for cultivar identification. Genetic similarity estimates derived from RAPD, simplified ^PstIAFLP and combined RAPD and ^PstIAFLP data were compared with coefficients of parentage (COP). Molecular markers-based mean genetic similarities were considerably greater than mean COP value. Correlation coefficients between COP and genetic similarities calculated from RAPD, ^PstIAFLP and combined molecular data were very low and not significant. A better correlation (0.50) was found between similarity estimates derived from RAPD and ^PstIAFLP markers. Four separate dendrograms were constructed based on pedigree and molecular analyses using a neighbor-joining algorithm (NJ). The dendrograms were compared and found to be topologically different. The results of this study showed, that both molecular techniques can be conveniently used for genetic characterization of oat cultivars, however ^PstIAFLP would be the method of choice due to the higher efficiency and reproducibility. This revised version was published online in August 2006 with corrections to the Cover Date.     

Changes in fatty acid composition associated with recurrent selection for groat-oil content in oat

Summary Groat oil of oat (Avena sativa L.) is a well-balanced oil with respect to saturated, monounsaturated, and polyunsaturated fatty acids. In this study, we measured the effect of six cycles of recurrent selection for high groat-oil content on the fatty acid composition of the groat oil. From each cycle of selection, 50 oat lines were grown in a replicated field experiment at two sites and were evaluated for groat-oil content and fatty acid composition.Concentrations of palmitate and linolenate decreased moderately over cycles of selection, whereas stearate content increased. A major increase occurred in the content of oleate and a major decrease in linoleate. Most of the modification of fatty acid composition took place within the saturated and unsaturated classes. The ratio of unsaturated to saturated fatty acids increased over cycles of selection. Significant genetic variation was present for each fatty acid, indicating that selection for different desired fatty acid compositions in groat oil of oat should be possible in this population.Abbreviations GO groat-oil content - 16:1 palmitate - 18:0 stearate - 18:1 oleate - 18:2 linoleate - 18:3 linolenate     

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.     

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

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

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.     

Effects of S1-recurrent selection for protein yield on seven agronomic traits of oat

Summary Three S₁-recurrent selection programs (designated HG, HP, and HGP) for increasing protein yield of oat (Avena sativa L.) were conducted for five cycles of selection. The selection criteria in each program (line of descent) emphasized different components of protein yield; high grain yield in HG, both high protein concentration of the groats (caryopsis) and high grain yield in HP, and high protein yield per se in HGP. Heading date and height were restricted to no net change. The objectives of this study were to evaluate the agronomic performance of these three lines of descent and the correlated responses in agronomic traits due to selection. Thirty to sixty random S₀-derived lines from each cycle and ten check lines were grown in hill-plot experiments at two locations in 1989. Grain yield increased in all three lines of descent; HG showed the greatest rate of gain followed by HGP and then HP (0.20, 0.10, and 0.07 Mg ha^-1 per cycle, respectively). Test weight decreased in HGP by 2.6 kg m^-3 per cycle and in HP by 4.0 kg m^-3 per cycle, but remained constant in HG. Cycle 5 (C₅) means for straw yield, height, and seed weight were not significantly different from the C₀ means for any line of descent. Groat percent remained unchanged in HGP, increased from 70.1 to 71.7% in HG, and decreased from 69.9 to 67.6% in HP. Heading date decreased by 3 d in HG and HGP, but remained unchanged in HP. HG compares most favorably with commercial cultivars because of its high yield and acceptable agronomic traits, while HP tends to be low in test weight, seed weight, and groat percent.Abbreviations BM- above ground biomass - GP- groat (caryopsis) percent, fraction of average seed weight that is groat rather than hulls - GPC- groat-protein concentration - GPY- groat-protein yield - GTY- groat yield - GY- grain yield - HD- heading date - HG- selection for protein yield through high grain yield - HGP- selection for protein yield per se - HP- selection for protein yield through high grain yield and high groat-protein concentration - HT- height - PY- protein yield - SDWT- seed weight - SY- straw yield - TWT- test weight     

Quantitative trait loci for quality and agronomic traits in two advanced backcross populations in oat (Avena sativa L.)

Using the advanced backcross quantitative trait loci (AB‐QTL) strategy, we successfully transferred and mapped valuable allelic variants from the high β‐glucan (BG) accession IAH611 (PI 502955), into the genome of cultivar ‘Iltis’. By backcrossing one BC₁F₁ plant to ‘Iltis’, we developed two BC₂F_2‐6 populations A and B, comprising 98 and 72 F₂‐individuals, respectively. Genotyping of BC₂F₂ individuals with predominantly AFLP markers resulted in 12 linkage groups with a map size of 455.4 cM for Population A and 11 linkage groups with a map size of 313.5 cM for Population B. Both populations were grown at three sites in Germany over a three‐year period. Individuals were then phenotyped for 13 traits including grain yield (YD) and β‐glucan content (BG). QTL analysis via stepwise regression detected a total of 33 QTLs, most of which were clustered in three linkage groups. Two dense linkage groups A1 and B13 were found to be putatively homologous to groups KO_6 and KO_11 of the ‘Kanota’/‘Ogle’ map, respectively.     

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.     

Direct and correlated responses to three S1-recurrent selection strategies for increasing protein yield in oat

Summary The objectives of this study were to evaluate the efficacy of S₁-recurrent selection for increasing groat (caryopsis)-protein yield in oat (Avena sativa L.) and to compare the effects of three selection strategies on correlated responses in groat yield and groat-protein concentration. Three S₁-recurrent selection programs were conducted for five cycles. Selection criteria in each program (line of descent) emphasized different components of protein yield; high grain yield in HG, both high grain yield and high groat-protein concentration in HP, and protein yield per se in HGP. Thirty to sixty random S₀-derived lines from each cycle and ten check lines were evaluated in one year at two locations to estimate the response to selection. Groat-protein yield increased 4.4% cycle^-1 in HG, 3.4% cycle^-1 in HP, and 2.2% cycle^-1 in HGP. Gains in groat yield were 7% cycle^-1 in HG, 1% cycle^-1 in HP, and 3% cycle^-1 in HGP. Mean groat-protein yield and groat yield in the cycle 5 (C5) populations of all three lines of descent were equal to or slightly higher than those of high yielding cultivars. The HG line of descent showed a significant decrease from 193 to 175 g kg^-1 groat protein, while the HP line of descent showed a significant increase from 202 to 218 g kg^-1. The HGP line of descent showed a small but significant decline in groat protein from 200 to 192 g kg^-1. In comparison, the high-protein commercial cultivars contained about 210 g kg^-1 groat protein. The C5 population of the HP line of descent had a unique combination of high groat yield and high groat-protein concentration. Genotypic variances for groat-protein yield, groat yield, and groat-protein concentration were not always significant in the various cycles of the three lines of descent, but in no instance was there a consistent change from C0 to C5. S₁-recurrent selection was effective for increasing groat-protein yield of oat, and with the proper germplasm and selection strategy both groat yield and groat-protein concentration can be improved simultaneously.Abbreviations C - cycle - GPC - groat-protein concentration (a groat is the part of an oat grain that is the actual seed or caryopsis, rather than the hull) - GPY - groat-protein yield - GTY - groat yield - HG - selection for protein yield through high grain yield - HGP - selection for protein yield per se - HP - selection for protein yield through high grain yield and high protein concentration Journal Paper No. J-15287 of the Iowa Agriculture and Home Economics Experiment Station, Ames, IA 50011, USA. Project No. 2447.     

Genetic divergence in two tropical maize composites after four cycles of reciprocal recurrent selection

Two tropical maize composites were subjected to four cycles of reciprocal recurrent selection to develop divergent inbred lines with good combining ability. This study was conducted to examine the extent of genetic diversity, changes in allele composition and genetic structure, of 100 randomly selected S₁ lines each from the original (C₀) and advanced (C₄) selection cycles of TZL COMP3 and TZL COMP4, genotyped using single nucleotide polymorphism (SNP) markers. Results revealed that the proportion of alleles at both low and high frequencies decreased from C₀ to C₄, whereas those at intermediate frequencies increased at C₄ in the two composites. More unique and other alleles were lost at C₄ in TZL COMP3 relative to those in TZL COMP4. The changes in different measures of genetic diversity were either small or negligible with selection in the two composites. The proportion of markers departing from Hardy–Weinberg equilibrium (HWE) decreased with selection, whereas the total number of pairs of markers in linkage disequilibrium increased with selection in the two composites. Examination of changes in population structures using a model‐based approach as well as cluster and multivariate analyses found a high degree of concordance in stratifying the 400 S₁ lines into four non‐overlapping groups corresponding to the two selection cycles each within the reciprocal composites. The observed molecular‐based divergence between cycles within the same composite and the clear differentiation between the complementary composites highlight the importance of reciprocal recurrent selection for preserving genetic diversity for long‐term selection. This increases the potential of the advanced selection cycles to sustain genetic gain in productivity of hybrids adapted to the savannas in West and Central Africa.     

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.     

Evaluation of strategies to optimize training populations for genomic prediction in oat (Avena sativa)

Genomic selection is a promising breeding methodology that could increase selection accuracy and intensity and reduce generation interval. As the cost of genotyping decreases, it will be important to optimize training populations for costly phenotypic experiments for many complex traits. The aim of this research was to evaluate different optimization strategies, by using historical data from the Norwegian oat breeding programme at Graminor. In this paper, we focus on the optimization criteria: genetic diversity, phenotypic variance and genetic similarity between the training and testing populations. The four training population strategies—prediction core, diversity core, phenotypic selection and random selection—were applied to an oat candidate population of 1124 lines. An independent testing population was used to calculate the mean prediction abilities for the traits days to heading and plant height. Moreover, the strategies were tested in three independent wheat populations. The results showed that prediction core was the most promising strategy to select training populations with high genetic similarity to the testing set, high genetic diversity, and high phenotypic variance, which resulted in higher prediction ability across population sizes and traits.     

A recurrent selection programme for grain yield in winter barley

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

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.