Evolution over 16 years in a bulk-hybrid population of subterranean clover (<Emphasis Type="Italic">Trifolium subterraneum</Emphasis> L.) at two contrasting sites in south-western Australia

Evolution was measured over 16 years in a self-regenerating, bulk-hybrid subterranean clover population, consisting of F ₂ seed from 253 crosses, sown at Nabawa and Mt Barker, short and long growing season sites, respectively, in south-western Australia. Seed bank samples harvested annually were grown in a common garden. Experiment 1 measured flowering time in plants from each year, while Experiment 2 measured 26 variables in the populations 3 and 16 years after sowing, in comparison with the ancestral population. Changes in population means were observed in 20 characters and variability declined in 11 characters at one or both sites, with much of this occurring within the first three years. Natural selection at Nabawa favoured early flowering of long duration, thick peduncles, high harvest index and high hardseededness. At Mt Barker it favoured late flowering of short duration, large leaves and long, thick petioles at flowering, thick stems with long internodes, long, thin peduncles with a high burial angle, large plants at maturity, low hardseededness and high biochanin A and total isoflavone contents. High seed production capacity, with high seed weight and seeds per burr, was important at both sites. The use of bulk-hybrid populations is demonstrated as a low-input means of breeding and selecting well-adapted subterranean clovers.

Structure and variation of subterranean clover populations from Sicily,Italy

Summary A collection of subterranean clover lines singled out from populations of the species Trifolium brachycalycinum and T. subterraneum collected in Sicily, Italy, was examined at two localities for flowering time and at one locality for oestrogen content and seed yield. The structure and variation of the populations of the two species were compared. The relationship between flowering time of the populations and some environmental features of their collection sites was examined to assess whether the maturity requirements of the two species were similar and to frame selection models focused on developing varieties of appropriate maturity. Populations of T. subterraneum were, on average, more complex than those of T. brachycalycinum, being characterized by higher number of lines per population and greater intra-population variation for flowering time, oestrogen content and seed yield. Furthermore, populations of T. subterraneum were, on average, about 15 days earlier than sympatric populations of T. brachycalycinum. Both the greater variation and the relative earliness of T. subterraneum occurred irrespective of the environments of origin of the populations. Inferences are drawn on the adaptive advantages that such features confer to T. subterraneum. Mean flowering time of the populations increased on increasing annual rainfall and altitude of the collection sites. However, the changes in maturity appeared almost exclusively related to variations in rainfall in T. subterraneum, while in T. brachycalycinum the effect of altitude was greater and that of rainfall less marked than in the former species.     

Influence of recurrent selection for flowering on flowering and yields in kura clover

Summary These investigations were conducted to determine the effectiveness of first- and second-season selection programs for increasing forage and seed yield in kura clover (Trifolium ambiguum Bieb.).Rhizo kura clover, the base population, was grown as spaced plants in two programs for six generations of selection for flowering in the seedling and second seasons. Remnant seed for each generation of selection was used to establish spaced plant and broadcast evaluation trials.The first-season selection program was effective in increasing flowering not only in the first season, but for two subsequent seasons. The second-season selection for flowering had little or no effect. However, vigor, forage and seed yields were not increased by either program, and the first-season program progressively reduced vigor and forage yields as generations of selection increased. The cause of this is unknown but may be associated with inbreeding depression or detrimental physiological effects of first-season flowering. Reselection to eliminate possible inbreeding effects is being conducted to further examine the possibility of increasing seedling and aftermath vigor in kura clover.     

Genetic variability and heritability estimates for hardseededness and flowering in balansa clover (Trifolium michelianum Savi) populations

Suitability of annual pasture legume species like balansa clover (Trifolium michelianum Savi) in southern Australian farming systems depends on their hardseededness and time to flowering. Late maturing varieties with increased rate of hardseed breakdown, particularly in the late summer—early autumn period are desirable to ensure reliable regeneration in permanent pastures. Two half-sib family balansa clover populations were used to estimate heritability for the rate of breakdown of hardseededness and time to flowering. High narrow-sense heritability estimates (90.5–96.1%, rate of hardseed breakdown; 85.9–94.5%, time to flowering) were obtained, and were mainly attributed to additive gene effects. There was no relationship between rate of hardseed breakdown and time to flowering in Group1 half-sib family, which indicated that selection for both traits could be undertaken independently. In half-sib family Group 2 the relationship between the two traits suggested the possibility of selecting late maturing cultivars with increased rate of hardseed breakdown. Significant differences in time to flowering and rate of hardseed breakdown both within and between the two half-sib family groups suggested that phenotypic recurrent selection would be effective in improving the traits in these populations.     

Six cycles of selection for adaptation in two exotic populations of maize

Summary The maturity rating of maize (Zea mays L.) grown in Galicia (Northwestern Spain) varies from FAO 200 to 600. Later germplasm has usually too much moisture content in the kernel at normal harvesting time so serious problems with storing could appear. Besides, farmers have tended to use earlier varieties in the last few years. All this imposes limitations on the amount of germplasm available to develop varieties adapted to this area.To study the possibility of adapting late, exotic material to the environment of the Atlantic coast of Galicia we carried out six cycles of individual selection on two non-adapted populations of maize (Purdue A and Purdue B). The criterion of selection was early silking and the criterion of response was moisture content of kernel at harvesting.The original populations and the populations obtained after each cycle of selection were crossed to the hybrids CM105 × CM109 (tester Reid) and H99 × H95 (tester Lancaster) and were evaluated in four environments to study the efficiency of the selection scheme. There were significant reductions in days to silking (6.1 and 6.7 days from cycle 0 to cycle 6 for Purdue A and Purdue B, respectively) and grain moisture at harvesting (3.0 and 3.9% from cycle 0 to cycle 6 for Purdue A and Purdue B, respectively). There were also reductions in plant height and yield in both populations.In general, the crosses population × Lancaster were higher for yield than the crosses population × Reid. Yield of the population crosses by both testers decreased after the six cycles of selection probably because of the earlier maturity of the selected populations. Some inbreeding depression may also have occurred.     

Breeding Strategies for Adaptation of Pearl Millet and Sorghum to Climate Variability and Change in West Africa

Semi‐arid and subhumid West Africa is characterized by high inter‐annual rainfall variability, with variable onset of the rainy season, somewhat more predictable endings, and drought or excess water occurrence at any time during the growing season. Climate change is predicted to increase this variability. This article summarizes options for plant breeders to enhance the adaptation of pearl millet (Pennisetum glaucum [L.] R. Br.) and sorghum (Sorghum bicolor [L.] Moench) to climate variability in West Africa. Developing variety types with high degrees of heterozygosity and genetic heterogeneity for adaptation traits helps achieving better individual and population buffering capacity. Traits that potentially enhance adaptive phenotypic plasticity or yield stability in variable climates include photoperiod‐sensitive flowering, plastic tillering, flooding tolerance, seedling heat tolerance and phosphorus efficiency. Farmer‐participatory dynamic gene pool management using broad‐based populations and diverse selection environments is useful to develop new diverse germplasm adapted to specific production constraints including climate variability. For sustainable productivity increase, improved cultivars should respond to farmer‐adoptable soil fertility management and water harvesting techniques. Larger‐scale, on‐farm participatory testing will enable assessments of varietal performance under evolving climatic variability, provide perspective on needs and opportunities and enhance adoption. Strengthening seed systems will be required to achieve sustainable impacts.     

Comparative studies within and between seed and seedling populations of leafed, ‘leafless’ and ‘semi-leafles’ genotypes of Pisum sativum L.

In an attempt to explain previously observed differences between the growth rates of leafless and conventional plants, we have compared representatives of three pea phenotypes; conventional (AfAfStSt), semileafless (afafStSt) and leafless (afafstst) with regard to their embryonic axis/embryo weight ratio and their patterns of pre-emergent growth. Within each phenotype, comparisons were made between genotypes which differed for mean weight per seed and between seed weight classes within each genotype. Although significant differences were found between genotypes for embryonic axis/embryo weight ratios, these could not be attributed to differences between phenotypes. Within each seed population there was a consistent decrease in this weight ratio with increasing seed size irrespective of phenotype.Relative growth rate (RGR) of the seedlings prior to emergence was similar for all genotypes within and between phenotypes and also for seed size classes from individual populations. There was however a positive relationship betweed seed size and absolute growth rate within and between seed populations.The results suggest that differences between the phenotypes do not become manifest until after emergence of the seedling from the soil and the possibility of a post-emergent lag phase in the leafless type is therefore suggested.     

Evaluating drought resistance in germplasm of meadow fescue

Summary The drought resistance of 25 accessions of meadow fescue (Festuca pratensis Huds.) from seven countries was investigated in four experiments: two in the glasshouse using pot-grown plants from which water was withheld for various periods, and two in controlled environments under osmotic stress.There were significant differences between populations (broad-sense heritabilities or repeatabilities) in all four experiments. In the glasshouse there was a large residual effect of yield potential on production during and after slight-to-moderate drought, and different susceptibilities appeared only after very severe drought. The most consistently high-yielding accessions were from the Bergamo alps in Italy. Recovery after drought was strongly correlated with tiller survival. Continued production under moderate drought was considered important in wetter climates, whereas survival and recovery under severe drought was often associated with low production or flowering in the seeding year, and more typical of summer-drought climates.Leaf growth rates of plants subjected to zero or moderate osmotic stress were correlated with yields of irrigated or moderately-stressed plants in the glasshouse, although there was no differential susceptibility to mild stress. Under severe osmotic stress there were very large differences in survival between populations, but there was no relationship with survival under glasshouse conditions.The inconsistency of population rankings across experiments shows that no one technique gives a full evaluation of drought resistance, and emphasizes the complex nature of the phenomenon. It was possible, however, to identify (a) regions meriting further collections, and (b) ecotypes that had desirable responses in all experiments and could contribute to variety improvement.     

Introgression of resistance to late blight (Phytophthora infestans) from Solanum pinnatisectum into S. tuberosum using embryo rescue and double pollination

In order to investigate the agricultural potential of the genus Vicia, and identify traits associated with productivity and responsiveness to environment, 34 undomesticated Mediterranean accessions representing Section Narbonensis (V. johannis, V. narbonensis) and V. sativa were grown in five contrasting environments in northern Syria (growing season rainfall: 76–290 mm).Highly significant genotype × environment interactions were observed for all traits. For most of the components of yield, accession mean performance (productivity)was highly correlated with responsiveness across environments (r = 0.59–0.96), as defined by joint linear regressions. Thus high yielding genotypes tended to be relatively more productive than low yielding genotypes under conditions that favoured high yields. Regression analysis revealed that mean site yields were positively correlated to rainfall (r = 0.85) and its attendant effect on growing season length as measured by cumulative season temperature and phenology (r = 0.59–0.81).In order to examine yield related traits independently of taxonomy, genotypes were grouped into three categories using K-means clustering based on productivity and responsiveness of seed, hay and biological yield. Highly productive/responsive genotypes were tall with high harvest index, large seeds and low fecundity (seeds and pods per plant), whereas unproductive/unresponsive plants tended to be short, highly fecund, with small seeds and low harvest index. Principal components analysis showed that responsiveness, in terms of seed, hay and biological yields, was closely related to phenological plasticity. Thus highly productive/responsive genotypes were able to start flowering earlier than unproductive/unresponsive genotypes in early environments, but significantly later in late, higher rainfall environments. Plant growth habit was also related to yield responsiveness. In environments with little biomass production the proportion of erect plants was high in all three categories. In more favourable, high biomass environments, the proportion of erect plants in unproductive/unresponsive genotypes fell dramatically, but was unchanged among productive/responsive genotypes. We suggest that for unproductive/unresponsive genotypes competition for light is increased under optimal growth conditions. We argue that the optimal combination of fixed and responsive traits in high yielding genotypes results in a `compound interest-type' response to more favourable environments. Highly productive and responsive genotypes can capture resources more effectively than their low yielding counterparts, leading to a positive relationship between performance and responsiveness for most components of yield. Differences in productivity and responsiveness for seed, hay and biological yield reflected Vicia taxonomy, increasing in the following order from low to high: V. johannis, V. sativa, the small seeded V. narbonensis (salmonea, jordanica, affinis) V. n. var. narbonensis, and finally V. n. var.aegyptiaca. V. n. var. aegyptiaca showed the most agricultural potential, since the taxon contained all the properties of productive/responsive genotypes listed above, yielding >1 t/ha under extremely arid conditions (104 mm),and >2.5 t/ha on 290 mm rainfall, confirming its potential for dry environments. This revised version was published online in August 2006 with corrections to the Cover Date.     

On defining a winter wheat

Summary An international survey of wheat breeders and scientists was conducted to determine whether there was a common understanding of the term winter wheat. Response to photoperiod was mentioned by only seven of the 30 survey respondents. It was consequently suggested that it be excluded from any definition. Response to vernalization was mentioned by 28 of the 30 respondents. However, there was considerable variation in the method of quantifying this response and the degree of response necessary for wheat to be called winter. It was concluded that the only objective and internationally consistent definition of winter wheat was a genetic definition based on the complement of Vrn genes a wheat possesses. Varieties not possessing any of the dominant Vrn alleles appear to take a distinctively longer and more variable time to head when grown under non-vernalizing, long day conditions. These are genetically winter wheats.     

Effect of Zinnia angustifolia HBK genotype on morphology and flowering of Z. angustifolia × Z. elegans Jacq. hybrids

Summary Interspecific hybrids were generated from crosses between 4 Zinnia angustifolia clones (maternal parents) and an inbred line of Z. elegans Orange King (paternal parent) to investigate the influence of Z. angustifolia genotype on morphology and flowering of hybrids. Leaf length, leaf width, flower diameter, number of ray petals, and days to flowering of interspecific hybrids were significantly influenced by Z. angustifolia clonal genotype. Genetic analysis of hybrid morphology and development was performed using 2 parental Z. angustifolia clones and 18 F₁ (9 seedlings from each of 2 populations generated from reciprocal crosses between parental clones) as maternal parents in crosses with Z. elegans Orange King. Genotype of Z. angustifolia significantly affected plant height, fresh weight, number of nodes, and days to flowering of interspecific hybrids. Control of these traits was through the Z. angustifolia nuclear genome. Improvement in interspecific hybrid morphology or development, i.e., shorter stature, earlier flowering, or larger flowers, may be possible by selection of superior-performing Z. angustifolia clones as parents.     

Effects of cytoplasm on heterosis and combining ability for agronomic traits in rice (Oryza sativa L.)

Summary Effect of sterilizing (WA) cytoplasm on heterosis and combining ability for days to flowering, plant height and grain yield in rice was studied in 70 crosses and their reciprocals produced by 10 cytoplasmic male sterile (A), their maintainer (B) and seven restorer (R) lines following line x tester design. The materials consisting of 140 hybrids (70 A/R and their reciprocal 70 R/B) plus 17 parental lines (10 B + 7 R) were evaluated under six environments, created by growing in three fields with different fertilizer doses (0, 60, 120 kg N/ha) and 2 seasons (dry and wet) during 1986 at IRRI farm. Reciprocal cross effect (A/R vs R/B) were highly significant for all the three traits. Interaction of reciprocal cross effects with environments were also highly significant for yield and days to flowering. Cytoplasmic effect for yield, days to flowering and plant height were estimated by comparing A/R and R/B combination and testing the significance of difference with LSD value. In order to avoid confounding effect of spikelet sterility on yields, twenty crosses showing normal spikelet fertility were selected. Both positive and negative cytoplasmic effects were observed for the three traits. The effects were modified by environments, except for plant height. Heterosis for all three traits was also affected by cytoplasm, however, manifestation of cytoplasmic effects was higher for heterosis for days to flower than in heterosis for yield and plant height. Effect of cytoplasm was more pronounced on general combining ability effects of parents than specific combining ability effects of crosses. Among the parents two CMS lines A4 (IR54752A) and A8 (IR22107-113-3-3A) and two R lines: R2 (IR46) and R7 (IR9761-19-1) showed consistent positive effect of cytoplasm on general combining ability. These lines have given several good heterotic combinations. The study indicated the usefulness of evaluating diverse cytoplasmic sources in various nuclear genotypes bred for hybrid rice breeding program.     

A new wilt-resistant parent for producing tomato rootstock seed

Summary Resistance to Verticillium wilt (VeVe) and Fusarium wilt, race 1 (II) has been bred into a genotype similar to that of GCR 66- a tomato rootstock seed parent used for the production of interspecific F₁ hybrid rootstock seed. The new seed parent is named GCR 115. It is proposed that the rootstock derived from GCR 115 be named Identistock KVF. The name Markerstock previously proposed in connection with GCR 66 has not been accepted by the registration authorities in the United Kingdom.     

Variation for dry matter yield,seed yield and other agronomic traits in Ladino white clover landraces and natural populations

Summary A set of Ladino white clover ecotypes including the last available landraces and some natural populations collected from old permanent meadows, together with some white clover varieties of various origins, were evaluated as monocultures for dry matter (DM) and seed yield assessed in separate field experiments. Other agronomic traits were also recorded. The best performing variety, Espanso, was significantly outyielded by some ecotypes for DM and seed production. Compared to landraces, natural populations yielded on average far less seed but as much DM and tended towards higher persistence indicating that domestication of the native Ladino at the farm level only improved seed yield of the crop. Natural populations showed, besides lower values of most seed yield components, a distinct canopy architecture characterized by smaller leaves, longer internodes and denser stolons. Seed weight proved the best character discriminating between the two ecotype groups. Significant variation for most traits was found both among landraces, in which it was mostly relatable to differences among provenances, and among natural populations. The relations between the main multivariate patterns of phenotypic variation and a set of environment collecting variables suggested that evolutive adaptation to different agronomic practices was a major determinant of the overall variation found in landraces. Evidence is provided that a relevant part of the variation observed among ecotypes for some important traits could be genetic.Abbreviations DM dry matter - DMY dry matter yield - NRH number of ripe heads - NFH number of florets per head - PL petiole length - LS leaf size - IL internode length - NS number of stolons - PRO protein content - SY seed yield - TSW thousand seed weight - NSF number of seeds per floret - PCA principal components analysis - PC principal component     

Genetic analysis of hexaploid wheat,Triticum aestivum using intervarietal chromosome substitution lines—protein content and grain weight

Summary The 21 intervarietal chromosome substitution lines of the cultivar Hope in Chinese Spring were used to analyse the genetic differences between the two cultivars Hope and Chinese Spring in grain protein content and grain weight.Only one chromosome of Hope, 5D, significantly influenced grain protein content of Chinese Spring. Its influence was of only minor effect and was to decrease protein content expression of Chinese Spring. It has been postulated that the genetic control of protein content, in this instance, is most likely due to many genes each of small effect.Five chromosomes of Hope influenced the 1000 grain weight value of normal Chinese Spring, all increasing its expression. Chromosomes 1A, 4A and 5B were of major effect and 3A and 6A of comparatively minor effect. A minimal estimate of five genes determines the difference in grain size between these cultivars. The possible evolutionary significance of the contribution of the A genome of bread wheat to grain size determination is discussed. On the basis of certain findings of this study, proposals are made for breeding for increased grain size in hexaploid wheat.     

Breeding for resistance to yellow disease of hyacinths. II. Influence of flowering time,leaf characters,stomata and chromosome number on the degree of resistance

Summary Analysis of an incomplete diallel design with 14 hyacinth cultivars and 4 wild accessions of Hyacinthus orientalis showed a significant GCA component for resistance to yellow disease. The late flowering hyacinth cultivars King of the Blues and Marconi and the early flowering Hyacinthus orientalis 70129 were found to be the best combiners for yellow disease resistance. Within a cultivar, the degree of earliness itself was not associated with the degree of resistance. Correlation coefficients of leaf characters and degree of resistance showed that hyacinths with short and/or narrow leaves are generally more resistant than those with long and/or broad leaves. Tetraploid cultivars with few, large stomata tend to be susceptible, diploid ones with many small stomata tend to be the more resistant.     

Backcross breeding to increase seed retention in cocksfoot (Dactylis glomerata L.)

Summary The objective was to evaluate the effectiveness of the backcross breeding method to improve an existing variety (Hallmark) for higher seed retention.The recurrent parent was cv. Hallmark (H) and the donor parent was cv. Marta (M). Marta is an Italian cultivar with a high degree of seed retention. Two cycles of backcrossing with selection for higher seed retention were conducted. In each cycle, 1200 plants were grown. In both BC₁ F₁ and BC₂ F₁, 20 plants were selected for general features of H and for a high degree of seed retention. The 20 plants of BC₂ F₁ were intercrossed in 1989. The bulked population thus obtained (HS) was evaluated in 1989–90 together with H and M as spaced plants in a randomized complete block design with 4 replicates. Regarding seed retention, H had a score of 7.9, M of 1.7 and HS of 2.8, LSD (0.05)=0.8, on a scale from 1 to 9. Consequently HS had similar seed retention to M but similar ear emergence date, plant height and panicle morphology to H. The results confirm that seed shattering is under control of a single partially or completely dominant gene.     

Resistance of wild emmer wheat to stem rust: Ecological,pathological and allozyme associations

Summary Seedling resistance to wheat stem rust was determined in populations of wild emmer wheat, Triticum dicoccoides, and characterized by means of ecological factors and allozyme genotypes. Reactions to wheat stem rust were studied in 102 single plant accessions of T. dicoccoides from ten populations by inoculation with Puccinia graminis tritici race 14, isolate GSR-739. Six populations displayed different degrees of response polymorphism with reactions ranging from high resistance to complete susceptibility, whereas four populations contained only susceptible plants. In some of the accession, unexplained intrasib variation in resistance and intraplant variation of infection-types were found. Resistance to stem rust was negatively correlated with two ecological factors, altitude and number of Sharav (hot-dry) days which are unfavorable to disease development. Variation in stem rust response was shown to exist in ecogeographic regions where climatic variables enhanced the development of the fungus, conceivably maintained by natural selection. Likewise, allozyme genotypes, single or in multiple loci combinations, appeared to be associated with resistance or susceptibility to rust. Such association need to be verified by genetic studies in order to become established as useful markers.     

Triple fusion of the primary endosperm nucleus as a cause of interspecific cross-incompatibility in Avena

Summary Using five diploid, two tetraploid and three hexaploid Avena species (x=7) 78 reciprocal crosses were made. Of these 74 were successfull, though the frequency of seed set varied greatly. The development of hybrid seeds differed widely between cross combinations, especially in reciprocal crosses. Hybrid seeds were grouped into four types, (1) normal kernels, well developed and germinative (designated as D+); (2) shrivelled-empty kernels, which do not germinate (E–); (3) small viable kernels (Rd+) and (4) small inviable kernels (Rd–). Some rare intermediate types were found in certain crosses.The results are well interpreted in terms of a hypothesis of polar-nuclei activation in which the strength of the activating action of the male nucleus, and the reaction of the female nucleus at double fertilization are expressed as activating value(AV) and response value(RV), respectively. The degree of seed failure is closely related to the activation index(AI) of the polar nuclei, AV/2RV (or × 100). In a selfed plant the activation index is always 1/2=0.5 (or50%), and this usually results in the formation of normal seeds. If AI deviates from 50% the endosperm often stops developing or degenerates.Based on the degree of abnormality of hybrid seeds, activating values of the ten species were arbitrarily assigned from 0.4 in A. ventricosa (2x) to 3.0 in A.sterilis (6x). For comparison A.strigosa (2x) was used as a standard and set at 1.0. Among the 74 succesful interspecific crosses the AI of the polar nuclei varied widely, from 7 to 375% Generally speaking, AI values of less than 20, 20–30, 30–80, and more than 80% show the R-, Rd+, D+ and E-kernel types, respectively. In other words, if the AV of the pollen parent is < 40 or >160% that of the maternal parent hybrid seeds fail to develop. Thus in double fertilization of Avena or probably angiosperms, the triple fusion of two polar nuclei with a secondary male nucleus is apparently a sexual isolation mechanism, being a barrier to interspecific hybridization.Professor Emeritus of Kyoto University, Faculty of Agriculture, Kyoto.     

Winter environment increases self seed yield in self-incompatible (SI) Petunia hybrida clones

Summary SI inbreds of P. hybrida and crosses between self-incompatible (SI) petunia plants were brought to flower under winter and summer glasshouse conditions. SI response, as measured by self seed, ranged from zero seed set under both conditions to low or zero seed set during the summer and high seed set during the winter. Some plants produced comparable seed yields during either pollination time. Increased expression of pseudo-self-compatibility (PSC) during the winter months was attributed to a breakdown of the Si system by the low light conditions of Minnesota winters (45°N). Genotypic differences within and between populations in the winter provided differences facilitating selection for increased SI. These differences were masked in some populations by the summer environmental conditions. The use of simulated or natural low light conditions as a technique for selecting for SI should increase selection efficiency.Scientific Journal Series Paper Number 10,499 of the Minnesota Agricultural Experiment Station.