Genetic analysis of drought stress response in rapeseed (Brassica campestris and B. napus). I. Assessment of environments for maximum selection response in grain yield

Summary Expectations of yield improvement in environments where drought was the major environmental factor limiting yields were studied in two species of rapeseed. Selection for yield in a drought stressed environment was predicted to be a more efficient selection stategy for yield improvement in dryland situations than selection in a more optimal environment, or selection based on a drought response index. The results indicate that selection for yield in a stressed environment are expected to lead to genetic advances in yield under optimal conditions as well as in a drought index. Selection under well watered conditions, on the other hand, was also expected to lead to correlated increases in yield in droughted environments but to decreases in the drought index. These results were found in both species of rapeseed grown in different water stress situations.The genetic advance in a drought response index was predicted to be greater in B. napus and marginally less in B. campestris if selection was practised for yield in a stressed environment rather than direct selection for the drought index. This was due to the higher heritability estimates in the stressed environments and the positive genetic correlations with yield.     

Genetic analysis of drought stress response in rapeseed (Brassica campestris and B. Napus). III. physiological characters

Summary Leaf proline accumulation, leaf chlorophyll stability, and germination at low osmotic potentials have previously been suggested as possible selection criteria for drought resistance in turnip rape (Brassica campestris) and rapeseed (B. napus) (Richards, 1978a). The feasibility of using these characters in a breeding program is reported. These characters were under significant genetic control and were responsive to selection. Broad sense heritabilities ranged from 40% for proline accumulation in both species to 55% for germination rate in B. campestris and 64% for chlorophyll stability in B. napus, however, narrow sense heritabilities for proline and germination were substantially lower. Their use as selection criteria for yield in a Mediteranean drought would be restricted since in B. campestris these characters were not related to yield while in B. napus only a weak association was observed between yield under drought and proline accumulation.     

Associations of some characters with seed yield in chickpea collections

Summary Three thousand two hundred and sixty-seven kabuli chickpea (Cicer arietinum L.) germplasm accessions were grown during the spring season of 1980 at Tel Hadya, the main research station of ICARDA, Syria to determine the components of seed yield. Observations were recorded on seed yield and 14 other characters. Correlation and path coefficient analyses were done to find out associations among characters and to assess the direct and indirect contribution of each character to seed yield.Large variation was observed for all the characters studied except days to flowering, days to maturity and protein content. Correlation and path coefficient analyses showed that biological yield and harvest index were the major direct contributors to seed yield. The 100-seed weight, plant height, days to flowering and maturity, canopy width, and protein content contributed to seed yield mainly through indirect effect via biological yield and harvest index. The 100-seed weight and seed yield were major contributors to biological yield. Major contributor to protein content was days to maturity. Results indicated that selection for high biological yield and harvest index would lead to high seed yield; and selection for large seed size would lead to high biological yield. Therefore, these characters should receive the highest priority in selecting high yielding plants in chickpea breeding.     

Selection for fast germination in rapeseed (Brassica napus L. and B. Campestris L.)

Summary Eleven populations of Brassica napus L. and twelve populations of B. campestris L. were subjected to three cycles of selection for fast germination at 2°C and at 25°C. The seeds from the selected populations, and unselected control populations grown in the same environment as the selected populations, were examined for germination behaviour at 2°C and 25°C, and for growth behaviour at 10°C. The populations in both species responded differently in terms of germination behaviour to selection for fast germination. In most of the populations that did respond positively to selection, selection practised at 2°C was superior to selection at 25°C in improving percent germination at 2°C, and was as good as the selection at 25°C in improving germination rate at the higher temperature. Selection for fast germination had no effect on growth characteristics of B. napus and B. campestris populations grown at 10°C. Thus, selection for fast germination at one low temperature may lead to improvement in germination characteristics over a range of temperatures, but will not necessarily lead to improved growth performance of the selected populations.     

Variation and association analysis in coriander

Summary Two hundred genotypes of Coriandrum sativum L. exhibited genetic variation for plant height, primary and effective branches, days to flowering and maturity, umbels and umbellets per plant, grains per umbellet, thousand seed weight, straw yield and grain yield per plant and harvest index. Heritability estimates were high for days to flowering, thousand seed weight and days to maturity; moderate for plant height, straw yield, umbels per plant, umbellets per plant and number of primary branches; and low for harvest index, effective branches, grain yield per plant and grains per umbellet. Phenotypic correlations of grain yield per plant were highly significant and positive with umbellets per plant, umbels per plant, number of effective branches, straw yield per plant, number of primary branches, plant height, number of grains per umbellet and harvest index. Maximum direct contribution to grain yield per plant was made by umbellets per plant, followed by straw yield per plant, umbels per plant and grains per umbellet. Umbellets per plant made sizeable indirect effect via straw yield per plant. Straw yield per plant made sizeable indirect contribution via umbellets per plant.     

Crossability and cytology of hybrid progenies in the cross between Brassica campestris and three wild relatives of B. oleracea,B. bourgeaui,B. cretica and B. montana

Summary Crossability and cytology were examined in F₁, F₂, B₁ and hybridsplants of F₁ hybrids of Brassica campestris and three wild relatives of B. oleracea, B. bourgeaui, B. cretica and B. montana, respectively. The F₂ plants were obtained after self-and open pollination of the F₁ hybrids. The B₁ and hybrid plants were produced after the F₁ hybrids backcrosses with B. campestris and crossed with B. napus, respectively. After crossing the F₁ hybrids, many seeds of the F₂, B₁ and hybrid plants were harvested. Multivalent formation was high in the chromsome configuration for the PMCs of F₂, B₁ and hybrid plants, suggesting that crossing over might occur between them. Many different types of aneuploids were obtained in the progenies of the F₂, B₁ and hybrid plants. It is suggested that different types of normal egg cells may be produced by one-by-one or little-by-little chromosome addition. The possibility is discussed of gene transfer from B. bourgeaui, B. cretica and B. montana, to cultivated plants, B. campestris and B. napus.     

Inheritance of seed colour in Brassica campestris L. and breeding for yellow-seeded B. napus L.

Summary Seed colour inheritance was studied in five yellow-seeded and one black-seeded B. campestris accessions. Diallel crosses between the yellow-seeded types indicated that the four var. yellow sarson accessions of Indian origin had the same genotype for seed colour but were different from the Swedish yellow-seeded breeding line. Black seed colour was dominant over yellow. The segregation patterns for seed colour in F₂ (Including reciprocals) and BC₁ (backcross of F₁ to the yellow-seeded parent) indicated that the black seed colour was conditioned by a single dominant gene. Seed colour was mainly controlled by the maternal genotype but influenced by the interplay between the maternal and endosperm and/or embryonic genotypes. For developing yellow-seeded B. napus genotypes, resynthesized B. napus lines containing genes for yellow seed (Chen et al., 1988) were crossed with B. napus of yellow/brown seeds, or with yellow-seeded B. carinata. Yellow-seeded F₂ plants were found in the crosses that involved the B. napus breeding line. However, this yellow-seeded character did not breed true up to F₄. Crosses between a yellow-seeded F₃ plant and a monogenomically controlled black-seeded B. napus line of resynthesized origin revealed that the black-seeded trait in the B. alboglabra genome was possibly governed by two independently dominant genes with duplicated effect. Crossability between the resynthesized B. napus lines as female and B. carinata as male was fairly high. The sterility of the F₁ plants prevented further breeding progress for developing yellow-seeded B. napus by this strategy.     

A study on disease variation in the populations of an interspecific cross of Brassica juncea L. x B. napus L.

Summary F₁ behaviour and F₂ variation in disease reaction were studied in the interspecific cross Brassica juncea x B. napus. Gene(s) for adult resistance to blackleg (Leptosphaeria maculans) were found to be present in the A genome of B. juncea and could be transferred to B. napus. Gene(s) for complete (seedling plus adult) resistance in B. juncea appeared to be located in the B genome. The chance of their transfer to the oilseed rapes (B. napus or B. campestris) would therefore seem to be remote.     

Transfer of resistance to race 2 of Plasmodiophora brassicae from Brassica napus to cabbage (B. oleracea var. capitata). II. Meiosis in the interspecific hybrids between B. napus and 2x and 4x cabbage

Summary Meiosis in 14 interspecific F₁ hybrids with three chromosomal levels (triploid, tetraploid, hexaploid; 2n=28, 37 and 55) between Brassica napus L. and 2x and 4x cabbage (B. oleracea var. capitata L.) was studied. The oleracea genome from B. napus maintained close homology with the c genome of cabbage while the campestris genome of B. napus showed partial homology with the c genome contained in the hybrids. Genotypic influence on chromosome pairing was indicated. Structural chromosome differences and spontaneous chromosome breakage and reunion were suggested as causes for the abnormalities which related to the unbalance of the genotypes. The divergence of the genomes of B. napus and B. oleracea and the need for the qualification of the term secondary association were discussed.Contribution No. J. 673, Research Station, Agriculture Canada, St. Jean, Québec.     

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.     

Associations between grain yield and carbon isotope discrimination in cowpea

Twenty cowpea (Vigna unguiculata L. Walp.) genotypes, comprising Kenyan cultivars and Kenyan and introduced breedings lines, were grown in 15 replicated field trials carried out at three locations in semi-arid eastern Kenya over four seasons. Grain and straw yields and the carbon isotope discrimination (Δ) of this material were determined, and days to flowering and maturity observed.None of the seasons in which the trials were conducted was appreciably drier than the long term average for the sites, and some trials received excessive rain. Averaging the data for the 15 trials, one genotype yielded much less grain (77 g m⁻²) than the 19 others, whose yield ranged from 125 to 177 g m⁻² (average 148 g m⁻²). Trial mean yield trials had a high straw Δ (r = +0.567) and had received more rain between flowering and maturity (r = +0.428) than lower yielding ones. Among genotypes, averaging over trials, the correlation coefficient (r) between grain yield and grain Δ was +0.394, and between grain yield and straw Δ, +0.460. Early genotypes had the highest grain Δ and straw Δ. The correlation among genotypes between date of flowering and grain Δ was −0.632 and with straw Δ, −0.502. When comparisons were made among trials, there was no clear relationship between the strength of the correlations among genotypes between grain yield and straw Δ or grain Δ and the degree of stress experienced by a trial (as indicated by the rainfall it had received). It appeared that this was because the variances of grain Δ and straw Δ were greater in the most droughted trials, whereas the variance of yield was greatest in the least droughted trials.It is concluded that the genetic correlation between grain yield and either grain Δ or straw Δ is unlikely to be strong enough to make either Δ a useful surrogate or adjunct in selecting for high yield in the segregating generations of a breeding programme for semi-arid eastern Kenya, but that Δ may be of value in the selection of parent genotypes.     

Assessment of drought resistance in 20 accessions of Triticum aestivum L. and related species II. Stability in yield components

Summary Stability in yield components was estimated in several species of Triticum grown across a range of soil moisture. The aim was to assess the effect of varying soil moisture on mechanisms leading to grain yield stability, particularly in the accessions identified as having stable yields and drought resistance viz., T. sphaerococcum, T. vavilovii and T. aestivum cv. C 306 (Paper I of this series). The study indicated that number of grains per spike provided stability to the accessions of T. sphaerococcum and T. vavilovii whereas number of spikes per unit area contributed towards the stability of T. aestivum cv. C 306. It is suggested that these accessions could be utilized as the genetic sources for improving specific yield components with regard to grain yield stability under droughted conditions.     

Interspecific transfer of Brassica juncea-type high blackleg resistance to Brassica napus

Summary Complete resistance to Leptosphaeria maculans, the cause of blackleg of oilseed rape (Brassica napus), was transferred from B. juncea to B. napus through an interspecific cross. B. juncea-type complete resistance (JR) was recognized first in one F₃ progeny (Onap^JR) by the absence of leaf-lesions on seedlings and canker-free adult plants. The commercially important characters of B. napus were retained in advanced lines of Onap^JR, which combined JR with low erucic acid levels (<0.5%), high seed yield and variable maturity dates.JR appeared to be inherited as a major gene or genes. Segregation for resistance and susceptibility contintied to occur during later generations of selection of Onap^JR. JR was readily transferred from Onap^JR to other suitable B. napus cultivars or lines with partial resistance to blackleg and resulted in highly vigorous carly generation selections adapted to cold, wet situations along with complete resistance to blackleg.     

The introgression of S alleles into forage rape,Brassica napus L. from turnip. Brassica campestris L. ssp. Rapifera

Summary Forage rape. B. napus, is self-compatible, the work described illustrates the introgression of functional incompatibility alleles into B. napus from turnips, B. campestris, in two generations. By grading seed, produced by backcrossing turnip/rape hybrids to rape, the frequency of 2n=38 semi articial B. napus recovered, amply justifies the cytological screening involved in such breeding programmes.     

Reaction of wild species of the geneus Lens to drought

Summary Susceptibility to drought stress is a key factor in dry land lentil (Lens culinaris Medikus) production in the Mediterranean region of West Asia and North Africa. This study examined the response to drought stress of 121 accessions representing all subspecies of the genus Lens; cultivated, and the wild L. culinaris ssp. orientalis (Boiss.) Ponert, L. culinaris ssp. odemensis (Ladiz.), L. nigricans M.B. Godr. ssp. nigricans Godr. and L. nigricans ssp. ervoides (Brign.) Ladiz. for their potential use in breeding for dry land conditions. Accessions were grown under two moisture regimes (dry land and dry land plus supplemental irrigation) at Breda, Syria during the 1990–91 and 1991–92 seasons. The cultivated lentil had markedly superior seed and straw production than did the wild Lens species. Time to flowering accounted for less than 10% of the variation in yield of wild accessions under rainfed conditions in the two seasons, showing that, in contrast to the cultivated germplasm, drought escape was relatively unimportant in wild lentil. Performance under drought in wild lentil, measured in terms of dry land seed yield or drought susceptibility index (S), was randomly distributed among collection locations with little relation to collection site aridity. Direct selection of wild lentil germplasm for biomass yield under dry conditions is of little value and an evaluation of wild accessions in hybrid combination is needed.     

Morpho-physiological traits to complement grain yield selection under semi-arid Mediterranean conditions in each of the durum wheat types mediterraneum typicum and syriacum

Summary Some 238 landraces from Algeria and Tunisia representative of the mediterraneum typicum durum wheat type and 265 landraces from Syria and Jordan representing the syriacum type were grown in a semi-arid Mediterranean environment of Northern Syria characterized by moderate drought stress. The germplasm types were compared for mean value, level of variation and relationships with grain yield of various morpho-physiological traits possibly usable for indirect selection of best yielding materials. The syriacum germplasm showed higher yield mainly due to greater earliness of cycle, slightly longer grain filling period, shorter stature, lower early growth vigour and higher drought tolerance expressed by a visual score recorded in another, more stressful environment in the region. It also showed lower variation for all morpho-physiological characters except plant glaucousness, for which it was more variable. The wheat types differed not only for architecture but also for optima of individual morpho-physiological traits required to maximize the yield response in the given environment. Higher yield of syriacum materials was attained at same heading and three to four days delayed maturity with respect to average phenology of three well-adapted control cultivars, and it was favoured by increasing number of kernels per spike, early vigour and drought tolerance. Higher yield of mediterraneum typicum landraces was related to heading and maturity dates approaching those of the control cultivars and to increasing kernel weight, early vigour and drought tolerance. Plant stature hardly affected the yield. Both absent and strong glaucousness could confer a yield advantage in syriacum materials.     

Implications of selecting improved strains of soybeans for dry-matter accumulation and grain yield

Summary Three maturity groups of soybeans (Glycine max L.) were used to investigate the relationship between dry-matter accumulation (DMA) and grain yield (GY), and the prospects for selection of high seed yielding strains among the existing soybean cultivars in a tropical environment. The positive and significant association between DMA and GY (r=0.888^***) indicated that selection for high DMA could give gains in GY. However, the higher harvest index (HI=37.5%) for the low seed yielding early maturing genotypes than the more vegetatively endowed and higher seed producing late maturity group, is an indication that excessive DMA could be disadvantageous. Total seed yield per land area for the three maturity groups of soybeans showed that the genotypes with high harvest index and low seed yield could be as good as those ones with high seed and dry-matter yields with low harvest index. The high coefficient of variation which ranged between 18.1 and 59.8% and the heritability estimations which also ranged from 34.4 to 82.2% are indicative of the presence of substantial genetic diversity and there are good prospects for the improvement of the crop through selection.     

Genetic effects on biomass yield in interspecific hybrids between Brassica napus and B. rapa

This study was conducted to estimate the genetic effects on biomass yield in the interspecific hybrids between Brassica napus and B. rapa, and to evaluate the relationship between parental genetic diversity and its effect on biomass yield of interspecific hybrids. Six cultivars and lines of oilseed B. napus and 20 cultivars of oilseed B. rapa from different regions of the world were chosen to produce interspecific hybrids using NC design II. Obvious genetic differences between B. rapa and B. napus were detected by RFLP. In addition, Chinese B. rapa and European B. rapa were shown genetically differences. Plant biomass yield from these interspecific hybrids were measured at the end of flowering period. Significant differences were detected among general combining ability (GCA) effects over two years and specific combining ability (SCA) effects differences were detected in 2000. The ratios of mean squares, (σ² _GCA(f) + σ² _GCA(m)) / (σ² _GCA(f) + σ² _GCA(m) + σ² _SCA), were 89% and 88% in 1999 and 2000, respectively. This indicates that both additive effects and non-additive effects contributed to the biomass yield of interspecific hybrids and the former played more important role. Some European B. rapa had significant negative GCA effects while many of Chinese B. rapa had significant positive GCA effects, indicating that Chinese B. rapa may be a valuable source for transferring favorable genes of biomass yield to B. napus. Significant positive correlation between parental genetic distance and biomass yield of interspecific hybrids implies that larger genetic distance results in higher biomass yield for the interspecific hybrids. A way to utilize interspecific heterosis for seed yield was discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Recurrent selection for yield in Glycine max using genetic male-sterility

Summary Recurrent selection, using genetic male sterility to facilitate intermating in Glycine max, was evaluated when selecting among individual space-planted S₀ soybean plants for three yield-related traits: 1) seed yield per se (YLD), 2) apparent harvest index (AHI), and 3) a selection index in which seed yield was regressed on maturity (REG). The original intermating population was a combination of the cultivar Century and two F₂ populations segregating for male sterility. The selection intensity through three cycles of selection was 20%. The YLD population increased in mean yield by 2.8±2.2 g plant^-1 cycle^-1 buf shifted significantly toward late maturity by 3.9±0.6 days cycle^-1. The AHI population decreased in yield by 4.4±2.6 g cycle^-1 while maturity shifted slightly earlier; there was very little change in AHI. The REG population increased in yield by 5.7±2.4 g plant^-1 cycle^-1 while shifting toward later maturity by only 0.8±0.5 day cycle^-1. We concluded that selection among S₀ plants in these populations, using the selection index of yield regressed on maturity, was affective in increasing single plant yields without causing a significant shift in maturity. Selecting for yield per se caused an unacceptable shift toward late maturity and selection for AHI was ineffective for increasing yield or AHI.     

Changes in morphological and physiological traits associated with recurrent selection for grain yield in maize

Summary Experiments were conducted during the growing seasons of 1975 and 1976 to determine changes in morphological and physiological traits associated with recurrent selection for grain yield in maize (Zea mays L.). Four variety hybrids, BSSS(R)CO x BSCB1(R)CO, BSSS(R)C7 x BSCB1(R)C7 [from a reciprocal recurrent selection program involving Iowa Stiff Stalk Synthetic (BSSS) and Iowa Corn Borer Synthetic #1 (BSCB1)], BS12CO x B14A, and BS12C6 x B14A [from a half-sib selection program involving the open-pollinated variety Alph (BS12) and the inbred tester B14A] were grown at 59 300 and 98 800 plants/ha near Ames, Iowa We obtained data on CO₂-exchange rate (an estimate of photosynthetic rate), grain yield, grain-yield components, flowering dates, maturity traits, light interception and use, shelling percentage, harvest index, and various other plant traits.CO₂-exchange rate did not change appreciably with recurrent selection for yield. Grain yield per hectare and per plant were larger for the improved than unimproved hybrids. Grain-yield components did not change significantly with recurrent selection. Kernel weight of BSSS(R)C7 x BSCB1 (R)C7, however, was larger than that of BSSS(R)C0 x BSCB1(R)C0. Pollen-shed-to-silking interval was shorter for the improved than the unimproved hybrids, and grain-filling duration was longer in C7 x C7 than in C₀ x C₀ of the reciprocal recurrent selection program. Furthermore, improved hybrids were characterized by smaller tassels and more upright canopies. Usually, plant traits and leaf-area-related traits were similar for all hybiids.Although dry-matter productivity was similar for all hybrids, those that were improved by recurrent selection produced more grain per unit leaf area and per unit light interception. Also, BS12C6 x B14A was characterized by a higher harvest index than BS12CO x B14A.We concluded that the source (i.e., photosynthetic capacity) was not limiting grain yield in BSSS(R) x BSCB1(R) and BS12. Increased grain yields that resulted from recurrent selection were consequences of longer grain-filling duration for BSSS(R) x BSCB1(R) and increased translocation of photosynthate from source to sink for both BSSS(R) x BSCB1(R) and BS12.Journal Paper J-8953 of the Iowa Agriculture and Home Economics Exp. Stn., Ames, Iowa. Project No. 2152.