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.     

施氮对青引1号燕麦生物量积累及其分配的影响

对‘青引1号’燕麦(Avena sativa‘Qingyin No.1’)采用不同施氮处理,探讨氮肥处理下各器官生物量积累和分配规律,以了解燕麦生物分配格局对氮肥响应机制,为燕麦合理施肥提供理论依据。结果表明:施氮显著影响了‘青引1号’燕麦各器官生物量积累和分配,在施氮75 kg/hm~2时茎、叶、穗、根和总生物量积累最高,分别达到1332.97、375.53、587.52、205.20、2501.22 g/m2;茎、叶、穗和根分配分别在30、60～90、90～120、0～60 kg/hm~2施氮处理下较高。‘青引1号’燕麦随生育期推进,总生物量、茎、根生物量呈先慢后快的增长变化,叶生物量呈先增后降的变化,穗生物量呈持续增加的变化。乳熟期生物量分配模式表现为茎穗叶根;茎、叶、穗和根生物量间均呈异速生长关系,茎、穗生物量的积累高于根,而叶生物量的积累则低于根。各器官在不施肥状态下的形态可塑性高于施肥处理。     

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.     

High Phytomass Producing Oats for Cultivation in Northern Growing Conditions

It has been argued that the breeding of cultivars with higher weight per panicle but constant phytomass (i.e., improved harvest index) is unlikely to improve grain yield of cereals in the future. To examine this question in oats, this study observed relationships between above-ground phytomass production and grain yield as well as 18 morphological and physiological traits representing maturity class and architecture of oat stand. The study was conducted in two dissimilar growing seasons. Material consisted of 29 Finnish and Scandinavian oat cultivars and breeding lines. Correlations between phytomass and other traits were evaluated by linear regression analysis. The study revealed that there has already been a trend toward improving the above-ground phytomass production in cereal breeding in Finland. The phytomass elevation can be attributed to both higher panicle weight as well as increased above-ground vegetative phytomass. Thus, improved grain yield of oats is not merely a consequence of increased harvest index, as is often mentioned, but is also a result of higher phytomass as such. The weight of the above-ground phytomass did not correlate with undesirable traits such as plant height or susceptibility to lodging, but it did have a positive relationship with days to yellow ripening. The applicability of phytomass weight in indirect yield breeding of oats in the northern growing conditions is discussed.     

Physiological Evaluation of Drought Resistance in Oats

Twelve varieties of Oat ( Avena sativa L.) were screened for relative drought tolerance based on the magnitude of chlorophyll stability index (CSI), relative water content (RWC) and free proline accumulation. IGFR1-2670, IGFRI-3021, IGFRI-2688 and IGO-8 showed a lower magnitude of CSI under stress condition in comparison to other tested varieties whereas the values of RWC and tree proline accumulation were higher. Based on the observations of this study it is concluded that, of the varieties tested, IGFRI-2670, 3021 2688 and IGO-8 are relatively more drought tolerant than the other genotypes tested. A close agreement was also noticed amongst the trends of CSI and proline accumulation, and per cent reduction in yield in rainfed over irrigated conditions.     

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.     

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.     

草燕1号燕麦新品种选育及品种特性评价

草燕1号燕麦（Avena sative‘Caoyan No.1’）是针对甘肃高寒地区及二阴地区旱作栽培条件下选育的高产、抗倒伏的燕麦新品种。该品种是以和政乡土燕麦为原始材料,采用单株混合选择法,经过近10年的选育而成。以青海444和青引1号为对照,在和政、定西、庆阳、天祝和夏河5个地点进行了品种比较试验、区域试验和生产试验。结果表明,草燕1号燕麦适宜在海拔1500~3000m的青藏高寒区和二阴冷凉区推广种植,其生育期比对照品种晚2~6d,牧草产量与对照无明显差异,但种子产量显著高于2个对照品种（P<0.05）,饲用品质优于青海444,且差异显著（P<0.05）,但与青引1号无显著差异。     

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.     

Konkurrenz und Ertragsvorteile in Gemengen aus Erbsen (Pisum sativum L.) und Hafer (Avena sativa L.)

In field trials on a fertile fluvisol in 1995 and 1996 near Göttingen, Germany, pea (Pisum sativum; cv. Messire/conventional leafed, cv. Profi/semileafless) and oats (Avena sativa; cv. Alf) were grown as sole crops and in substitutive mixtures. The sole crops were established at 80 pea seeds m^?2 and 300 oat seeds m^?2. The mixtures consisted of 67 % (pea) and 33 % (oats) of the monoculture densities, respectively. Interactions of cv. Messire or cv. Profi and oats were similar in 1995 and 1996. The mixtures outyielded the monocultures with respect to total above ground dry matter (RYT = 1.15) and grain yield (RYT = 1.09). Grain yield of pea and oats averaged 4.9 t ha^?1 in monocultures and 5.5 t ha^?1 in mixtures. Oats was relatively the stronger of the two competitors. Decreasing number of pods per plant could be highlighted as the factor for a lower pea seed frequency in the yield of the mixtures. For oats the number of panicles per plant and kernels per panicle were higher in the mixtures compared with the oat monocultures. The average amount of the harvest index (HI) was 0.52 for pea and 0.46 for oats. Favourable growth conditions increased HI values however, prolific vegetative growth in the mixtures resulted in lower HI values. The predicted RYT‐values estimate the maximum combined grain yield of 6.3 t ha^?1 in the mixture of 87 % pea (70 seeds m^?2) and 13 % oats (39 kernels m^?2).     

Expression of nuclear-cytoplasmic interactions and heterosis in quantitative traits of oats (Avena spp.)

Summary Nuclear and cytoplasmic genes from 10 diverse Avena sterilis L. accessions were introgressed into four Corn Belt oat (A. sativa L.) cultivars. Grain and straw yield, harvest index, heading date, height, unit straw weight, and vegetative growth index were evaluated in the BC₂ generation of 76 cytoplasmic isopopulations. Means of all seven traits in the BC₂ of Tippecanoe matings were greater than Tippecanoe due to introgression of A. sterilis germplasm, whereas means of all seven traits in BC₂ populations of Ogle were inferior to those of Ogle. BC₂ populations of matings involving CI 9170 and CI 9268 were variable, with some trait means being greater than those of the recurrent parents and some being lower.The phenomena of nuclear-cytoplasmic interaction effects and nuclear-cytoplasmic heterosis also were investigated. In matings where significant differences were present between cytoplasmic isopopulations, those with A. sterilis cytoplasm usually were superior, but no trait exhibited a consistent cytoplasmic effect across all matings. These results suggest that all seven traits were influenced by nuclear-cytoplasmic interactions. Nuclear-cytoplasmic heterosis for grain yield was present in two of 38 isopopulations with A. sterilis cytoplasm; both were from matings involving Tippecanoe. Nuclear-cytoplasmic heterosis for each of the remaining traits was observed in four to eight of the 38 isopopulations with A. sterilis cytoplasm.     

Responses in Agronomic Traits Associated with Three Recurrent Selection Regimes for Groat-Oil Content in Oat*

Crop strains with high values for the selected trait but unacceptable agronomic characteristics are not useful as varieties. In this study, we determined whether selection for groat-oil content in oat (Avena sativa L.) via three recurrent selection regimes caused associated changes in 10 agronomic traits. Random lines of each cycle of selection of each regime were evaluated in a replicated field experiment at two locations. Biomass, grain yield, groat yield, and test weight decreased in all three regimes. Seed weight and plant height decreased in two regimes, and groat fraction in one. Oil yield increased in two regimes and decreased in the other. Plants headed earlier in one and later in another regime over cycles of selection. No changes in harvest index occurred. Heritabilities and genetic variances were relatively high for all traits in each regime. Factor analysis of the phenotypic correlation matrices indicated that simultaneous increases in grain yield and in groat-oil content are possible, that continuous selection for groat-oil content decreases groat fraction and test weight, and that seed weight is strongly and negatively associated with groat-oil content. A new recurrent selection regime for increasing groat-oil content of oat that may overcome the negative responses of agronomic traits was proposed.     

Vererbung des β-Glucangehaltes der Haferkaryopse

Inheritance process for β-glucan content in oat caryopsis
Due to the fact that there are only few studies concerning β-glucans in oat caryopsis with regard to plant-breeding aspects, three oat crosses were performed (cross 19 = low × high, cross 86 = low × low, cross 98 = high × high). In all of the three oat crosses, lower β-glucan content (% DM) was recorded in F, than in F, indicating that the β-glucan content (% DM) at a higher heterozygoty grade is lower than at a reduced heterozygoty grade. This points to a dominant inheritance of lower β-glucan content (% DM). A tendency towards the female β-glucan content (% DM) could not be proven. Thus, it can be started from a nuclear genetic inheritance of the β-glucan content (% DM). In most cases, the experiments showed a normal distribution of β-glucan content (% DM), indicating for several genes a quantitative inheritance of β-glucan content (% DM). By means of the regression of the F₃-lines to the F₂-plants, the narrow-sense heritability could be calculated for the cross 86 (h²= 0.51***) and the cross 98 (h²= 0.48***). The heritability for the cross 19 was not significant.     

Effect of Water Stress on Biochemical and Physiological Characteristics of Oat Genotypes

Activities of enzymes catalase, peroxidase, and level of total soluble proteins were studied in two genotypes (JHO 884, JHO 888) of oat ( Avena sativa L.) subjected to water stress for 1, 3, and 4 days by withholding water in a pot culture experiment. No change was observed in the level of total soluble proteins at early stage but it increased significantly at higher magnitude of water stress. Increase in catalase activity was noticed in JHO 884 whereas reverse trend was observed in JHO 888 at highest level of stress. The JHO 884 genotype showed increase in peroxidase activity from the first day of water stress. Results suggest that the efficiency level of antioxidant system is found different in these two genotypes.     

Leaf Area Duration of Oat at High Latitudes

Contribution of leaf area duration (LAD) to grain yield during the short growing season characteristic of northern latitudes may differ from the marked impact it has at lower latitudes. Three experiments (exps) were carried out at Viikki Experimental Farm, University of Helsinki, Finland (60° 13′N) to compare associations between main shoot and tiller LAD with grain yield, yield components and morpho-physiological traits characterizing plant stand structure. This was done using correlation analysis and principal component analysis (PCA) for data from trials conducted during 1993–1995. Exp I included three N fertilizer rates (80, 120, 160 kg N ha^?1), three seeding rates (250, 500, and 750 seeds m^?2), and tall, moderate, semi-dwarf, and dwarf oat lines. Exp II incorporated two N fertilizer rates (80 and 120 kg ha^?1), three seeding rates (400, 600, and 800 seeds m^?2), and naked and hulled oat lines, and exp III, foliar applications of plant growth regulators [control, chlormequat chloride (CCC), and ethephon], and dwarf, naked, and conventional oat lines. LAD for main shoots and tillers [calculated as ∫ LAI d T, where T is cumulated degree days from seedling emergence to yellow ripening (dd °C)] and 12 morpho-physiological traits were measured. Oat was not able to benefit from high main shoot LAD if drought occurred at grain-fill, but in the absence of severe drought or if it occurred at pre-anthesis, high LAD favoured yield formation. Tiller LAD had a negative impact, if any, on grain yield in cases of an inverse relationship between tiller LAD and yield components on main shoots was recorded. Use of low seeding rates resulted in improved ability of total LAD to contribute to dry-matter production, but was not associated with grain yield. Low seeding rates enhanced formation of high tiller LAD, which was not able to compensate for grain yield reduction caused by fewer main shoots. Foliar application of CCC and ethephon tended to increase the proportion of LAD production by tillers at the expense of grain yield, which can be attributed to lower post-anthesis precipitation. Oat types differed in main shoot and tiller LAD. Disadvantageous characteristics of naked oat, such as low single groat weight, were not associated with insufficient main shoot LAD. High tiller LAD in naked lines and the Minnesota-adapted dwarf line that was not associated with high tiller grain yield in naked lines, in particular, indicates that tiller growth was enhanced at the expense of grain yield production. Application of N fertilizer at various rates did not have any effect on LAD.     

灌溉条件下燕麦氮、磷、钾配方施肥效应分析及与产量回归模型的建立

为探讨灌溉条件下燕麦品种银燕6号氮（N）、磷（P）、钾（K）配方施肥的肥料效应,采用“3414”试验方案,研究N、P、K不同施用配比对燕麦产量和农艺性状的影响,建立了施肥与产量的回归模型,为构建灌溉条件下燕麦施肥指标体系提供科学依据。结果表明,处理N₂P₂K₂籽粒产量（5100.0kg/hm²）、肥料贡献率（34.8%）、产值（20 400.0元/hm²）在14个处理中均最高;增产效应为N>P>K,并且N、P、K肥间存在明显的交互作用;N、P、K单种肥料施用量与籽粒产量呈抛物线关系,根据拟合函数得出,N、P、K最大施用量分别为229.8、80.5、26.4kg/hm²;根据N、P₂O₅和K₂O施用量对籽粒产量的影响,建立了三因素的施肥数学模型,进一步得出,燕麦产量≥4950.0kg/hm²时,氮（N）︰磷（P₂O₅）︰钾（K₂O）施肥比例为1.54︰1︰0.23,N、P₂O₅、K₂O施用量范围分别是148.8~198.3、>112.5和14.3~37.2kg/hm²。该结论对灌溉条件下燕麦生产具有实际指导意义。