Climatic adaptation in subterranean clover populations

Summary Long-term persistence and, hence, agronomic success as a pasture of the annual species subterranean clover depend primarily on seed yield and seed survival over seasons. In natural populations, plant characteristics influencing seed setting and formation of seed reserves in the soil are expected to be adjusted to the prevailing environmental conditions of the sites of origin. Knowledge on plant/environment relationships may provide information on adaptive strategies of persistence, and guidelines for selecting adapted varieties to specific conditions. On pure lines from a number of populations such relationships were assessed for flowering time, seed yield, burr fertility, individual seed weight, initial hardseededness, and rate of hardseededness breakdown over summer. Flowering time decreased on decreasing annual rainfall, i.e., on shortening the growing season, as adaptive response to the need of producing adequate seed before the onset of the dry season. Individual seed weight decreased on decreasing rainfall, and increasing temperatures. Hard-seed maintenance over summer was higher in populations from hot and dry environments, where the marked effect of temperature on hardseededness breakdown exerts a strong selective pressure. Within-population variation, assessed on flowering time, was particularly wide, with early genotypes occurring even in populations from long-season environments. The adaptive relevance of maintaining high levels of within-population polymorphism to cope with unpredictable climatic fluctuations is discussed. Number of constituent lines as a measure of the population structure, and intra-population variation were both influenced by altitude and rainfall, tending to decrease as the climatic selective pressure becomes severe, under both low-rainfall, hot conditions and high-elevation, cold-prone environments.     

Geneticanalysis of drought stress response in rapeseed (Brassica campestris and B. Napus). II. Yield improvement and the application of selection indices

Summary The expected improvement in grain yield if selection was made for plant characters measured between flowering and maturity was evaluated in populations of rapeseed (Brassica campestris and B. napus) grown in a droughted environment. Drought was commenced at flowering in each species and measurements were made on plants at the commencement of the drought stress, during the stress treatment and at crop maturity.Substantial genetic and phenotypic variation was observed in yield as well as the different morphophysiological determinants of yield. In B. campestris no single parameter was found to be a suitable alternative selection criterion to yield since the correlated responses in yield if selection was for another character was lower than if selection was for yield alone. By the use of a selection index however, joint selection for yield, as well as harvest index, 1000 seed weight and seeds per pod, was expected to be 20% more effective than direct selection for yield under drought. In the B. napus population direct selection for flowering time or for harvest index was predicted to result in a genetic advance in yield equal to or greater than that obtained by direct selection for yield, whereas joint selection for flowering time and yield should result in a 16% greater yield increase. Selection for increased yield in these populations grown in droughted conditions is discussed.     

Geographical patterns of morphological variation in sorghum (Sorghum bicolor (L.) Moench) germplasm from Ethiopia and Eritrea: Quantitative characters

A total of 415 sorghum (Sorghum bicolor (L.) Moench) accessions representing different regions of Ethiopia, Eritrea and a group of introduced lines were evaluated for 15 quantitative characters to determine the extent and geographical pattern of morphological variation. The extent of variation was highly pronounced for agronomically important characters for sorghum. These characters included plant height, days for 50% flowering, peduncle exsertion, panicle length and width, number and length of primary branches per panicle and thousand seed weight. Significant regional variation was also observed for most of the characters. The results implied that environmental factors such as altitude, rainfall, temperature and growing period are important in regional variation. Mean for plant height and for days for 50% flowering showed clinal variation along the gradients of rainfall pattern and growing period in Ethiopia. Moreover, there were significant positive correlation coefficients between most of the characters. This included the correlation between agronomic characters of primary interest in sorghum breeding such as plant height and days for 50% flowering and also between various characters and the altitude of the collection sites. The implications of the results in plant breeding, germplasm collection and conservation as well as the probable sources of the wide range of variation are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Some yield components and ideotype traits in meadowfoam,a new industrial oil crop

Summary Three populations of Limnanthes alba and five of L. douglasii were investigated for relative growth rates, leaf area index, flowering time, seed and oil yield, seed size and shattering. Within each species the early flowering populations had the highest seed yield. Most accessions developed adequate leaf canopy by the time of onset of flowering.An ideotype for breeding goals could be defined in terms of the characters of L. douglasii var. nivea such as early flowering and high seed yield, provided taller plant stature and reduced seed shattering are incorporated.Domestication and breeding might involve a combination of induced mutation, interspecific hybridization and conventional breeding procedures.     

Variation for reproductive and agronomic traits among T. repens × T. nigrescens third generation backcross hybrids in the field

Interspecific hybridisation with Trifolium nigrescens Viv. is a possible strategy to improve the reproductive potential of white clover (Trifolium repens L.). Following the development of a fertile F1 hybrid, three generations of backcrossing have been carried out usingT. repens as the recurrent parent. Vegetative characteristics, stolon growth and seed yield components of the backcross (BC) 2 and 3 generations, as well as the parental species were measured on spaced plants grown in the field. Leaf size and plant spread of the BC2 and BC3 generations were less than T. repens but there was no difference in plant fresh weight. Numbers of inflorescences per plant and florets per inflorescence of the backcrosses were greater than T. repens however this was not reflected indifferences in seed yield per plant asT. repens had more seeds per floret and per plant than the backcrosses. Differences in stolon length, the proportion of flowering nodes and the pattern of axillary bud development were observed between T. repens and the backcrosses. Significant variation among the BC 3 generation for vegetative and reproductive traits was observed. Individual plants among the BC 3 generation were identified that combine high forage yield, substantial inflorescence production and good fertility, and these will form the basis of further selection. This revised version was published online in August 2006 with corrections to the Cover Date.     

Variation between and within lentil landraces from Yemen Arab Republic

Summary The variation in 31 landraces of lentil collected from Yemen Arab Republic was partitioned to estimate the variation between populations and the variation between families within populations by studying a total of 568 progenies of randomly selected plants. There was significant variation between populations and between families within populations in seedling pigmentation, time to flower, time to maturity and plant height, but the predominant source of variation was between populations. This variation was unrelated to altitude of collection. The landraces were equally variable across characters. The implications of this analysis of variation on lentil breeding and germplasm collection are discussed.     

Trifolium ambiguum as a source of variation in the breeding of white clover

Summary Hybrids between T. ambiguum and T. repens were successfully established after ovule culture. The hybrids were male sterile but one of the hybrids produced a single seed when backcrossed to T. repens, while the other produced a significantly greater number of seeds. The majority of the backcross hybrids from the latter had 48 chromosomes, being produced from the fertilisation of unreduced gametes of the F1 hybrid by T. repens pollen. Regular meiosis in the BC1 hybrid resulted in the production of balanced gametes containing the full complement of T. repens chromosomes and the polyhaploid set of T. ambiguum chromosomes. There was no difficulty in establishing a substantial population of BC2 hybrids. The meiotic behaviour of the BC1 hybrid indicated that it should be possible to transfer genes from T. ambiguum into white clover T. repens. The significance of the results in terms of the use of alien variation in the improvement of white clover is discussed.     

Genetic variation for seed dormancy in Cuphea laminuligera and Cuphea lanceolata

Summary Various Cuphea species are adapted to temperate climates and characterized by caprylic, capric, and lauric acid-rich seed oils. These fatty acids are solely commercially available from palm kernel and coconut oils; thus, there is considerable interest in Cuphea as an alternative source of these fatty acids. Cuphea is a genus of undomesticated species characterized by seed dormancy and seed shedding. Our goal is to eliminate these domestication barriers in a few species with agronomic potential. Our objective was to survey existing Cuphea laminuligera and C. lanceolata germplasm for genetic variation for seed dormancy (germination percentage after a certain seed storage period). Recurrent half-sib family selection was practiced in various open-pollinated and synthetic populations. Half-sib family seed were evaluated for germination percentages at 26° C after various seed storage periods. There was significant genetic variation among families in every population. Additive genetic variance, half-sib family-mean heritability, and expected selection response magnitudes depended on the length of the seed storage period within a given population. Estimates of these parameters were significantly greater for longer seed storage periods within a population. Heritability estimates ranged from 0.00 to 0.79. Expected half-sib family selection responses ranged from 0.0 to 23.9%. Our study demonstrated there is substantial genetic variation for seed dormancy in C. laminuligera and C. lanceolata.     

Quantitative trait loci for early maturity and their potential in breeding for earliness in Brassica juncea

A doubled haploid population of Brassica juncea, developed from a cross between two parental lines differing for days to maturity, was used to study the efficiency of indirect selection for a primary trait through selection of secondary trait(s) over direct selection for the primary trait when quantitative trait loci information is available for both primary and secondary traits, and applied. Days to maturity was considered as primary trait, while days to first flowering, days to end of flowering, flowering period and plant height were considered as secondary traits. An RFLP linkage map was employed for QTL analysis of maturity and maturity-determinant traits, and a stable QTL B6 simultaneously affecting these two types of traits was identified. This linked QTL explained 11.7% phenotypic variation for days to maturity, 20.7% variation for days to first flowering, 24.3% variation for days to end of flowering and 14.4% variation for plant height. Phenotypic evaluation of maturity and/or maturity-determinant traits, viz. days to first flowering, days to end of flowering and plant height revealed that limited genetic advance for early maturity can be achieved through phenotypic selection of the primary and/or the secondary trait(s). However, the estimates of genetic advance for early maturity based on combined phenotypic evaluation and linked QTL data was found to be, at least, three times higher compared to genetic advance based on phenotypic evaluation only, demonstrating the potential of marker-assisted selection in breeding for early maturity in B. juncea.     

Genetic control of flowering time and its component processes in subterranean clover (Trifolium subterraneum L.)

Summary The inheritance of flowering time and its component processes, vernalization and photoperiod response, were studied in two crosses of subterranean clover (Trifolium subterraneum L.) using a field sowing and four controlled environment sowings with different combinations of vernalization and photoperiod. Time to flowering was under polygenic control and was highly heritable. For both vernalization and photoperiod response, there was dominance for a low response, or earliness. A simple genetic control was indicated for photoperiod response. The results for vernalization response were not clear cut, although the character appeared to be under polygenic control. An interaction between vernalization and photoperiod response was evident in three of the four cultivars studied. This made it impossible to separate the effect of these two component processes and complicated the study of their inheritance. Node of first flower on the main stem was closely related to flowering time and its use led to similar conclusions in the inheritance studies.     

Combining ability for seed yield traits of Chenopodium quinoa breeding lines

For a better understanding of the inheritance of seed yield traits in Quinoa (Chenopodium quinoa) Willd., a half-diallelic crossing experiment with six diverse but uniform breeding lines was conducted. True hybrid plants were detected by means of differences in panicle colour in 14 out of the 15 crosses performed. The agronomic performance of 14 F₂ populations and six parental lines was evaluated in field trials on fertile clay soil. General and specific combining ability of the lines (GCA and SCA) were estimated. Highly significant differences in GCA effects were found between the lines for plant height at maturity, early flowering, early maturity, seed yield and thousand seed weight. Significant SCA effects were only found for plant height at maturity. However, SCA effects were rather small and accounted for 7% of the total variance. The finding of large GCA effects and low SCA effects suggests that best selection results could be expected from crosses between the agronomic best performing genotypes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic variation in a Norwegian germplasm collection of white clover (Trifolium repens L.)

Genetic variation and genotype × environment interactions for important agronomic characters were estimated for eleven Norwegian populations of white clover (Trifolium repens L.), originating from between 58^°52′N and 69^°30′N latitude, and from altitudes of 10 to 450 m a.s.l. The populations were evaluated in monoculture clonal field plots for two years at two locations in Norway. The medium-leaved Danish cultivar Milkanova was also included in the experiment as a standard market cultivar. Considerable variation between populations was observed for winter survival, spring growth, morphological characteristics, dry matter yield, general performance, earliness and seed yield. Winter survival, measured as plant cover after two years, was significantly different at the two locations. All the populations had sufficient winter survival at the southern, continental highland site (61^°07′N).At the northern coastal location (69^°30′N),where the winter climate is unstable with physical stresses such as frost, ice-encasement and water-logging frequently occurring, only the northernmost, coastal populations showed sufficient winter survival. Highly significant interactions were found between populations and locations, and between populations and years for all characters measured at both locations and in both years, respectively. Interaction effects between populations, locations and years were significant only for leaflet length. The largest leaf lamina's were observed in the most southern populations. However, genotypes possessing relatively long leaflets were also found in the northernmost populations, although at low frequencies. In general, the populations originating from the northernmost locations had lower biomass production, and were smaller-leaved, more prostrate and exhibited shorter internodes and stolons compared to the populations from South and Middle Norway. These populations also initiated inflorescences earliest, and were among the lowest seed yielding populations. Otherwise, no clear clinal variation in growth characteristics was observed which could be related to the latitude or altitude of origin of the populations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Morphological variation in Astragalus hamosus L. and Coronilla scorpioides L. populations of Tunisia

Morphological variation between Astragalus hamosus and Coronillascorpioides populations was studied using local germoplasm collected in northern and central Tunisia. Twenty-one morphological traits were recorded and data were analysed using complementary statistical analysis. Considerable variation based on morphological and agronomical traits was found between populations and for both species. Pod and peduncle lengths as well as flowers number were the most discriminant variables between populations. Differences were also found for variables like seed number per pod, seed yield and dry matter production. Significant correlations were found between plant traits and the environmental parameters of the origin sites. This study revealed enough variation among populations to initiate a selection programme for pasture improvement in arid and semi-arid areas of Tunisia, where the two species are well adapted. This revised version was published online in July 2006 with corrections to the Cover Date.     

Intra-varietal variation for in vitro plant regeneration in the genus Trifolium

Summary Seedlings of Trifolium repens showed considerable variation with regard to the morphology and growth of their calli, and their ability for in vitro differentiation of shoots. One of the lines selected for regeneration in primary callus cultures also showed shoot formation from protoplasts. Somatic embryogenesis in callus cultures of T. pratense and T. arvense occurred only in selected seedling lines. This paper highlights the importance of screening a large number of plants within a cultivar of outbreeding species to achieve reproducible plant regeneration from tissue culture.     

Relationship between morphological variation and geographical origin or selection history in Lupinus pilosus

Patterns of morphological diversity were examined in Lupinus pilosus in relation to geographical origins or selection history of accessions. There was significant variation among accessions for most architectural and reproductive characters in a field experiment in Perth, Western Australia. Late flowering was associated with resetted early growth, profuse branching and collection in higher rainfall sites. High yield and profuse podding on the main stem was associated with large pods and leaves, many seeds per pod, greater height to the first main-stem pod, and collection in higher altitude sites. Ten groups of accessions, identified by hierarchical cluster analysis, accounted for 79% of genotype and 69% of genotype x character sums of squares. One group, possibly taxonomically distinct from the others, consisted exclusively of wild types from northern Israel with poor pod set and low seed yield, poor nodulation, pale foliage, and small leaves. Wild types from Turkey and Crete had rosetted early growth and were late flowering, and the Turkish group had extremely rapid mid-season growth. Ornamental types from Europe or Australia with pink, purple or white flowers clustered together in an early flowering group with vigorous early growth. A smooth-seeded type of L. pilosus was discovered in a group of short, late flowering and rough-seeded types from Syria.     

Genetic diversity of wheat wild relatives in the Near East detected by AFLP

In order to reveal the molecular genetic diversity of wheat wild relatives, an AFLP analysis was conducted with 16 accessions of five Triticum andAegilops species originating from the Near East. Variation within population was studied with at least seven individuals per accession. Four primer combinations were used for selective amplification. Based on the scored bands, we estimated percentage of polymorphic bands, 1 – proportion of shared bands (1-psb) and nucleotide diversity (π). Of the five species used in this study, Ae. speltoides had the highest level of `within population' variation. This species had also the highest value of the variation among populations. As for Triticum species, the level of variation within population was low in diploid species (T. urartu and T. boeoticum),whereas two tetraploid species (T. dicoccoides and T. araraticum) had relatively high levels of variation within population. While the two diploid Triticum indicated a clear interspecific divergence, the two tetraploid wild wheats were not clearly divergent in this study. The variance portioning analysis indicated that the variation detected for diploid Triticum species was mainly composed of `between species' variation, on the other hand that for tetraploid Triticum was mostly composed of `within population' variation. In conclusion, AFLP analysis reveals molecular variation in all accessions used in this study, suggesting a potential genetic diversity of the wheat wild relatives in natural populations. These results have implications for the design of strategies to maintain genetic diversity within genebank collections. This revised version was published online in August 2006 with corrections to the Cover Date.     

Polymorphism of natural populations of Pinus halepensis Mill. in Morocco as revealed by morphological characters

In Morocco, Aleppo pine (Pinus halepensis Mill.) is the most used species in reforestation. However, no data is available about its intraspecific variation and adaptability. Morphological traits of 15 natural populations located on a North-East South-West orientation, representing its natural distribution in Morocco, were used to assess geographic variation of this species. Seed size (length, width, depth, weight and 1000 seed weight), wing length and width, cone length and width and needle length were measured for 760 trees (43–65 trees per population). A correlation matrix between morphological characters and geographic parameters (altitude, latitude north and longitude west) exhibited negative and positive correlations of seed size with the latitude and longitude parameters, respectively. Also cone characters correlated positively with altitude. Thus, populations originating from the South-West region (High Atlas and a part of Middle Atlas) showed high values of Aleppo pine characters. Factorial component analysis (FCA) and hierarchical classification of all populations lead to the formation of two major groups (South-West and North-East) containing, each one, two minor groups. Geographic pattern of Aleppo pine distribution and its adaptability in Morocco are discussed in this study. This revised version was published online in July 2006 with corrections to the Cover Date.     

Characterization of flowering time and SSR marker analysis of spring and winter type Brassica napus L. germplasm

Flowering dates and life forms of all available Brassica napus accessions conserved at the North Central Regional Plant Introduction Station (NCRPIS) were characterized, and a survey of molecular variation was conducted by using simple sequence repeats (SSR) in order to support better management of accessions with diverse life forms. To characterize flowering phenology, 598 B. napus accessions from the NCRPIS collection were planted in Iowa and Kansas field sites together with a current commercial cultivar and observed for days to flowering (first, 50% and 100% flowering) in 2003. Days from planting to 50% flowering ranged from 34 to 83 in Iowa and from 53 to 89 in Kansas. The mean accumulated growing degree days (GDD) to 50% flowering were 1,997 in Iowa, and 2,106 in Kansas. Between locations, the correlation in flowering time (r = 0.42) and the correlation in computed GDD (r = 0.40) were both significant. Differences in flowering-time rank were observed for several accessions. Accessions that failed to flower in Iowa in a single growing season comprised 28.5% of the accessions; of the flowering accessions, 100% plant flowering was not always achieved. Accessions were grouped according to flowering time. A stratified sample of 50 accessions was selected from these groups, including 10 non-flowering and 40 flowering accessions of diverse geographic origins and phenological variation. The flowering time observed in the sampled accessions when grown in the greenhouse were found to be significantly correlated to the flowering time observed in the field locations in Iowa (r = 0.79) and Kansas (r = 0.49). Thirty SSR markers, selected across 18 Brassica linkage groups from BrassicaDB, and 3 derived from Brassica expressed sequence tags (ESTs) were scored in the stratified sample. An average of three bands per SSR primer pair was observed. Associations of SSR marker fragments with the life forms were determined. Analysis of molecular variation by using cluster analysis and ordination resulted in recognizable, distinct groups of annual and biennial life-form types, which may have direct applications for planning and management of future seed regenerations. Mention of commercial brand names in this paper does not constitute an endorsement of any product by the U.S. Department of Agriculture or cooperating agencies.     

Breeding the common bean (Phaseolus vulgaris L.) for resistance to bean common mosaic virus: alternatives to backcrossing

A series of field experiments was undertaken in order to determine whether resistance to bean common mosaic virus (BCMV) could be incorporated into genotypes of the common bean (Phaseolus vulgaris L.) suitable for cultivation in Zimbabwe without recourse to backcrossing. Six inbred genotypes carrying the resistance-conferring alleles at the loci I and Bc-3 were crossed with five locally-adapted inbred genotypes. The first experiment comprised F₃ progeny rows, each derived from a single unselected F₂ plant, the second, F₃ bulks selected for resistance, and the third, a comparison of selected and unselected F₂-derived F₄ lines. The number of days to flowering and to maturity, the incidence of mosaic and necrosis symptoms, seed yield and seed size were recorded. There was evidence that late flowering and maturity were associated with BCMV resistance in some crosses, though not strongly enough to present an obstacle to plant breeding. The incidence of virus symptoms and seed yield were influenced by genetic factors additional to the major resistance genes, and variation in seed yield was present not only between bulk populations of crosses, but also between single-row plots of lines within crosses. This indicates that early-generation selection for yield in the presence of BCMV, even among progeny selected for BCMV-resistace, is likely to be effective. However, the variation in yield among F₄ lines was least in the highest-yielding crosses, which may represent a limit to successful selection for yield. Seed size was partly under additive genetic control, but there was also evidence of non-allelic interactions. There was no association between large seed size, preferred by consumers, and susceptibility to BCMV in the progeny, indicating that the association between these characters in the parent lines is fortuitous and will not present an obstacle to plant breeding. It is noted that a considerable amount of useful genetic information can be obtained without recourse to elaborate crossing schemes, provided that unselected progeny are included in experiments as controls. The evidence presented indicates that resistance to BCMV can be combined with appropriate values of maturity date, yield and seed size without the need for backcrossing.