The presence or absence of the soybean Kunitz trypsin inhibitor as a quantitative trait locus for seed protein content

Numerous quantitative trait loci (QTL) for various characters have recently been reported in different crop plants. However, information is limited about the molecular mechanisms behind QTL, because most of them have only been detected at a statistical level. Therefore, progeny from a cross between two soybean genotypes segregating for the presence vs. absence of the Kunitz trypsin inhibitor, a 21.5 kDa protein, have been analysed for possible effects of that protein on agronomic and seed quality characters. Protein content was reduced by, on average, 4.5 g/kg in segregants lacking the Kunitz protein, whereas oil content and other characters remained unaffected. This finding can be interpreted as a ‘model QTL’ for variation in seed protein content, because the molecular and genetic backgrounds of the soybean Kunitz trypsin inhibitor are well understood.     

Impact of low linolenic acid content on seed yield of winter oilseed rape (Brassica napus L.)

An essential quality improvement of rapeseed oil can be obtained by reduction of its linolenic acid (C18:3) content from about 10% to less than 3% of the total fatty acids. Genotypes low in C18:3 have been developed by mutagenesis. The initial summer rapeseed mutant had been low yielding and highly susceptible to various diseases. It has been debated whether the low C18:3 character can be successfully combined with high seed yield for physiological reasons. Therefore, the low linolenic character of mutant M48 was transferred into high-yielding genotypes by repeated backcrossing to well-adapted low erucic acid, low glucosinolate (00-) winter rapeseed cultivars. Lines with low C18:3 content were selected from BC3 and BC4 generations and examined in 1990–95. Positive selection response for seed yield was shown to continue over the years. Presently, the best lines are yielding as well as the control cultivars being equivalent also in oil and glucosinolate contents. In order to test the effect of a low C18:3 content on seed yield, plants with low and with high C18:3 content, respectively, were selected from 16 segregating BC5-F₂ populations and bulked to form 32 F₃ populations. These ‘isogenic’ bulk populations were tested for field performance at four locations in 1995. The results show that C18:3 content of the seed oil is not associated with seed yield, oil content, beginning of flowering, plant height and disease resistance. Means of relative seed yield for the high and the low linolenic F₃ bulk populations were not significantly different with 88.0% and 86.9% of the control cultivars, respectively. There was a significant interaction between genotypes with high or low C18:3 content and location. This shows that under specific environmental conditions a low C18:3 content may be either favourable or unfavourable. The results indicate that the low C18:3 character of the original mutants per se does not cause a decrease in seed yield, oil content or general field performance.     

Identification of quantitative trait loci underlying seed protein and oil contents of soybean across multi‐genetic backgrounds and environments

Seed protein and oil contents are important quantitative traits in soybean. Previously, quantitative trait loci (QTL) associated with seed protein and oil were mostly identified in single genetic background. The objective of this work was to identify QTL and their epistatic effects underlying seed protein and oil contents in three recombinant inbred line populations (two of them used one common female parent) across eight environments by composite interval mapping. Forty QTL underlying protein content and 35 QTL underlying oil content were identified. Among them, nine were universal QTL underlying protein content and four were universal QTL underlying oil content. Epistatic interactions between QTL underlying seed protein/oil and different genetic backgrounds were detected. Different pairs of epistatic interactions were observed in diverse genetic backgrounds across multi‐environments. Common marker intervals were observed to simultaneously underlie seed protein and oil contents with different epistatic interactions. The results in this study suggested that a specific genotype with high oil content and low protein content might significantly affect the selection of soybean lines for high seed protein.     

Alternative strategies of recurrent selection for seed yield of soybean

Summary Recurrent selection programs can be manipulated by varying either the extensiveness of testing used to identify superior lines or the number of intermating generations conducted among the superior lines between cycles of selection. The research was conducted to compare the performance of soybean (Glycine max (L.) Merr.) lines derived from populations developed by recurrent selection for seed yield using a factorial arrangement that combined one or two stages of replicated yield testing to identify superior lines with one or three generations of intermating among the selected lines. The base population AP6, which was used for this evaluation of alternative strategies of recurrent selection, was derived from 40 high-yielding strains of Maturity Groups 0 to IV. For this study, the 40 original parents of AP6 and the lines used as the parents for the most advanced cycle of selections for each of the strategies were evaluated in four Iowa environments. The number of cycles of selection completed for the strategies under evaluation varied from two to five. Strategies with two stages of replicated yield testing to select superior lines resulted in a larger number of high-yielding lines than when only one stage of testing was conducted. A larger number of high-yielding lines was obtained from strategies with one generation of intermating than when three generations were used. With the procedures used for the one-stage and two-stage tests in this study, the results indicated that an appropriate strategy for recurrent selection of seed yield in soybean would involve two stages of replicated yield testing to identify superior lines and one generation of intermating among the selected lines to form a new population.Journal Paper no. J-12025 of the Iowa Agriculture and Home Economics Experiment Station, Projekt 2475, Ames, IA.     

Environmental effects on lutein content and relationship of lutein and other seed components in soybean

Lutein is a major carotenoid in soybean [ Glycine max (L.) Merr.] seed, and has been shown to be beneficial for eye health in humans. Development of soybeans high in lutein is a goal of some breeding programmes. Little is known about how different growing environments affect lutein content. The objective of this study was to determine the variation of lutein and its relationship to seed protein, oil and individual fatty acids in soybean seed. Fifteen soybean genotypes were planted at four environments. There was no effect of year and planting date on lutein content in soybean seed. However, genotype × year, genotype × planting date and genotype × year × planting date were significant for lutein content. Although each genotype showed similar response across environments, lutein content varied significantly across the four growing environments in 14 of the 15 genotypes evaluated. Lutein content was not correlated with seed protein or oil and palmitic or stearic acid concentrations. However, lutein was positively correlated with oleic acid content and negatively correlated with linoleic and linolenic acids content.     

Seed quality attributes of food-grade soybeans from the U.S. and Asia

Diversity of food-grade soybeans is critical for utilization of genetic resources in cultivar development, germplasm enhancement, and end-product commercialization. The objective of this study was to assess seed quality attributes and phenotypic variability among 54 U.S. and 51 Asian food-grade cultivars and breeding lines. The results showed greater genetic diversity of protein content, calcium content, seed hardness, and seed size uniformity than other quality traits in both small- and large-seeded genotypes evaluated in this study. Among the small-seeded soybeans, the U.S. genotypes were more diverse and exhibited higher swell ratio and oil content but lower stone seed ratio and protein content than Asian accessions. Among the large-seeded accessions, U.S. genotypes had higher stone seed ratio and oil content but lower swell ratio and protein content, and were less diverse than Asian genotypes. The characterization of diverse food-grade soybeans will facilitate parent selection in specialty soybean breeding.     

High Protein Soybean Performance Under Field Conditions

An early maturing soybean [ Glycine max (L.) Merr.] cultivar has been developed with substantially higher seed protein concentration than previously available cultivars. The objectives of this study were to evaluate the performance of the high protein line under field conditions in Eastern Canada, and to investigate the interrelationships between protein and oil concentrations, seed weight and yield. A field experiment, including the high protein line OT89-16 and its recurrent backcross parent Maple Arrow grown side by side, was conducted at McGill University, during 1990 and 1991. The protein concentration of OT89-16 was 17 % greater than that of Maple Arrow for both years. However, the protein yield of OT89-16 was not significantly increased, the oil content was 19 to 23 % lower, and oil yield per ha was over 30 % lower than those of Maple Arrow. The lower yield of OT89-16 was closely associated with its lighter seeds. Further efforts to improve the high protein line should attempt to maintain the same high protein levels but achieve yield equal to that of the parent line through selection for heavier seeds.     

Selection for protein content in soybean from single F<Subscript>2</Subscript> seed by near infrared reflectance spectroscopy

Soybean seed averages about 40% protein and is a dominant source of protein in animals and human foods. Increasing protein in soybeans is a goal in soybean breeding programs. Initial selection for protein among progeny from breeding populations is often based on non-destructive Near Infrared Reflectance (NIR) spectroscopy analysis of a 5 g or more seed sample in the F₃ or later generations. NIR spectroscopy analysis for protein on single F₂ seed would allow selection at the earliest possible time and improve selection efficiency for protein content; however this practice has not been evaluated. The objective of this study was to test effectiveness of NIR spectroscopy analysis of single F₂ seeds to select for increased protein content from two populations developed from high and average protein content parents. F₂ seeds from a single F₁ plant of each population showed a normal distribution with transgressive segregation for protein concentration. In both populations, F₃ seeds produced from plants from single F₂ seeds with either low or high protein content were analyzed for protein by NIR spectroscopy. Protein means and ranges of F₃ seed selected from high protein F₂ seeds were higher in protein than F₃ seed from low protein F₂ seeds which produced low means and ranges in protein content. This shows that analysis of single F₂ seed for protein content using NIR spectroscopy was effective in selecting for increased protein in the F₃ generation. Analysis of single F₂ seeds from breeding populations will improve breeding efficiency for protein in soybean breeding programs.     

Characterization of genotype by environment interactions in soybean breeding programmes of southeast Europe

Southeast Europe is Europe's major production area of soybean of maturity groups (MG) 0, I and II, but genotype by environment interaction (GEI) for soybean traits in this region is still not characterized. The objective of this study was to characterize GEI in soybean grown in Croatia for seed yield, protein and oil content. Fourteen soybean cultivars of MG 0 and I were evaluated at 15 environments. In the combined anova , genotypes, environments and GEI were significant. All three seed traits were more affected by year than by location and seed yield and protein content were more sensitive to environmental changes than oil content. Genotype by environment interaction is generally of less importance than effects of genotypes and years, especially for oil content. High‐yielding genotypes had values of regression coefficient (b) close to 1, indicating that they were not responsive to extreme environments, either positive or negative. Means and b values were not correlated for yield and negatively correlated for protein and oil content, probably because of low‐input practice in the region. The absence of recognizable differences in means, b values and principal component scores of ammi analysis between two MGs for all traits indicated that soybean cultivars of MG 0 may be recommended for growing in the region because they responded similarly to environmental changes as do full‐season genotypes of MG I.     

Predicting transgressive segregants in early generation using single seed descent method-derived <Emphasis Type="Italic">micro-macrosperma</Emphasis> genepool of lentil (<Emphasis Type="Italic">Lens culinaris</Emphasis> Medikus)

In a self-fertilised crop like lentil, the identification of transgressive segregants for economically important trait such as seed yield is an important aspect of any practical breeding programme. The prediction of expected transgressive segregants in F₁ generation obtained as a ratio of additive genic effect [d] and additive variance (D) i.e. [d]/√D was studied in 28 crosses of lentil generated in a diallel fashion involving four parents each of macrosperma (exotic) and microsperma (Indian) types, respectively, resulting in three hybridization groups. The seed material advanced to F₂, F₃ and F₄ generations through single seed descent method was evaluated to determine the observed transgressive segregants for seed yield/plant. The observed frequency of crosses showing more than 20% transgressive segregants in F₂ to F₄ generations were exhibited in 9(32%) crosses, of which 7(77%) crosses were of macrosperma × microsperma type. Genotypes Precoz and HPL-5 of the exotic group (macrosperma) produced maximum number of transgressive segregants with the genotypes L-259, L-4145 and PL-406 of the Indian origin (microsperma). Goodness of fit (non-significant χ² value) in F₂ generation was observed for 19(68%) crosses of the total genepool, out of which 9(56%) crosses each in F₃ and F₄ generation belonged to the macrosperma × microsperma group, depicting it as the gene pool of paramount importance to obtain maximum transgressive segregants, therefore establishing the efficacy of the method used.     

Dissection of genetic architecture for oil content in soybean seed using two backcross populations

Soybean seed oil was valued in foods, animal feed and some industrial applications. Molecular marker‐assisted selection (MAS) for high‐oil‐content cultivars was an important method for soybean breeders. The objective of this study was to identify quantitative trait loci (QTL) and epistatic QTL underlying the seed oil content of soybeans across two backcross (BC) populations (with one common male parent ‘Dongnong47’) and two different environments. Two molecular genetic maps were constructed. They encompassed 1046.8 cM [with an average distance of 6.75 cM in the ‘Dongnong47’  ×  ‘Jiyu89’ (DJ) population] and 846.10 cM [with an average distance of 5.76 cM in the ‘Dongnong47’  ×  ‘Zaoshu18’ (DZ) population]. Nine and seven QTL were identified to be associated with oil content in the DJ and DZ populations, respectively. The phenotypic variation explained by most of the QTL was usually less than 10%. Among the identified QTL, those stable ones across multiple environments and populations often had stronger additive effects. In addition, three stable QTL in the DZ populations were identified in the similar genomic region of the three QTL in the DJ population [qDJE and qDZE‐1 were located near Satt151 of Chromosome 15 (Chr15), qDJA1 and qDZA1 were located near Satt200 of Chr15 (LG A1), and qDJD2‐1 and qDZD2‐1 were located near Sat365 of Chr17]. In conclusion, MAS will be able more effectively to combine beneficial alleles of the different donors to design new genotypes with higher soybean seed oil content using the BC populations.     

Selecting recombinants to stack high protein with high oleic acid and low linoleic acid in soybean (Glycine max)

Soybean accounts for over a quarter of the world's oilseed consumption and over 70% of the world's protein meal consumption. The separate development of high oleic, low linolenic acid (HOLL) soybean and high-protein (HP) soybean means that no soybean cultivar on the market has an optimal fatty acid profile and increased protein. The objective of this study was to develop and evaluate high protein, high oleic acid, and low linolenic acid (HP-HOLL) soybean. A five-gene stack was created using a two-phase forward breeding scheme and marker-assisted selection method. Forty-six HP-HOLL lines from three genetic backgrounds were grown in six environments in the Southeast United States. Although genotype-by-environment interaction was significant for seed composition traits, lines met the >75% and <3% cutoffs for oleic acid and linolenic acid, respectively, and met or exceeded the protein concentration of the HP parent. No negative interaction could be detected between the HP and HOLL traits. Additionally, yield testing in four environments indicated yield parity for some lines, suggesting HP and HOLL soybean cultivars with high yield could be selected.     

Effects of high oleic acid soybean on seed yield,protein and oil contents,and seed germination revealed by near‐isogeneic lines

Typical soybean oil is composed of palmitic, stearic, oleic, linoleic and linolenic acids. High oleic acid content in soybean seed is a key compositional trait that improves oxidative stability and increases oil functionality and shelf life. Using a marker‐assisted selection method, near‐isogenic lines (NILs) of G00‐3213 for the high oleic trait were developed and yield tested. These NILs have various combinations of FAD2‐1A and FAD2‐1B alleles that were derived from the same backcrossing populations. The results indicated that G00‐3213 NILs with both homozygous mutant FAD2‐1A and FAD2‐1B alleles produced an average of 788 g/kg oleic acid content. The results also demonstrated that possessing these mutant alleles did not cause a yield reduction. Furthermore, seed germination tests across 12 temperatures (12.8–32.0°C) showed that modified seed composition for oleic acid in general did not have a major impact on seed germination. However, there was a possible reduction in seed germination vigour when high oleic seeds are planted in cold soil. The mutant FAD2‐1A and FAD2‐1B alleles did not hinder either seed or plant development.     

Mapping quantitative trait loci controlling soybean seed starch content in an interspecific cross of ‘Williams 82’ (Glycine max) and ‘PI 366121’ (Glycine soja)

Seed starch content (SSC) greatly affects the taste, flavour and processing properties of soy foods. The objective in this study was to identify quantitative trait loci (QTL) for SSC in soybean. A total in 169 recombinant inbred lines (RILs) derived from a cross in ‘Williams 82’ and ‘PI 366121’ were grown for three consecutive years. The SSC of the RILs displayed continuous variation with transgressive segregation and hence amenable for QTL mapping. Nine significant QTL exhibiting 5.6–11.3% of the total phenotypic variation (PVE) were identified. The QTL qSTR06_2 showed highest PVE (9.1–11.3%) at LOD values of 4.25–5.39. No stable QTL over 3 years were identified, indicating strong environmental influence on SSC. The QTL qSTR11_1 and qSTR20_1 were found to colocalize with some of the previously reported QTL for sucrose content in soybean, implying the interrelationship between starch and sucrose biosynthesis. As the carbohydrate components may affect key constituents such as oil and protein in soybean seed, findings of the study may be useful in breeding soybeans with improved seed composition.     

Comparison of pedigree selection and single seed descent for oil yield in linseed (Linum usitatissimum L.)

Summary Four populations of linseed derived from five parents were advanced from the F₂ to the F₆ generation by SSD and to the F₅ by pedigree selection. These populations were used to compare the efficiency of the two methods in order to provide superior genotypes with respect to grain yield and oil content, i.e. oil yield. The results showed minimum differences between pedigree and SSD lines for grain yield, where in only one cross the SSD lines were significantly superior to the pedigree lines. Since pedigree selection was carried out for both, seed yield and oil content, a positive response to selection was expected. However, early selection for yield, a character with low heritability, was not successful. On the contrary, significant differences for oil content were detected between the two groups of lines in three of the four crosses studied. In these cases the pedigree lines were superior to the SSD lines. These results demonstrate that strict selection in early generations for oil content, a character with comparatively high heritability, is feasible and successful in linseed. However, selection for seed yield should be postponed to later inbred generations. Consequently, in breeding for maximum oil yield of linseed a two-step selection procedure is recommended.Abbreviations PS pedigree selection - SSD single seed descent - TGW thousand grain weight     

Generation and characterisation of colchicine-induced polyploid Lavandula × intermedia

Double podding in cultivated chickpeas (Cicer arietinum L.) can increase yield and yield stability. In the present study, we performed reciprocal crosses of ‘kabuli’ (double podded) and ‘desi’ (single podded) chickpeas to determine (i) the expressivity and penetrance of double podding, (ii) the correlations of yield and yield components, and (iii) the heritability of double podding, flower color, and stem pigmentation in F₂ plants. Reciprocal crosses were performed with two genotypes, AC 2969 (kabuli) and ICC 4969 (desi), to generate F₁ and F₂ plants. The results indicated hybrid vigor (heterosis) for yield in F₁ plants and better performance of F₂ plants. Yield and yield components of some lines in F₂ were superior to the best parent, indicative of transgressive segregation. In particular, the presence of double podding (‘s’ allele) significantly increased yield in some of the transgressive segregants. Expressivity and penetrance of the ‘s’ allele depends on the background of the female parent. Some of the double podding progeny had greater seed yields than those of the single podding progeny and greater seed yields than the best parents. Double podding, stem pigmentation, and pink flowers each appears to be governed by a single recessive gene. Stem pigmentation and pink flowers appear to be linked traits that depend on the genetic background of the crossed chickpeas. Taken together, our studies of reciprocal crosses of kabuli and desi chickpeas clearly showed that yield could be improved by selection for transgressive phenotypes that have double podding.     

Effects of divergent selection for seed protein content in high‐protein vs. food‐grade populations of early maturity soybean

Due to the growing need for vegetable protein in Central European agriculture, there is interest in producing food‐grade soybeans, which are higher in seed protein and sucrose content and have a larger seed size than conventional soybeans. As protein content of conventional soybean is often below 400 g/kg, either high‐protein or food‐grade donors were crossed with adapted genotypes in order to increase their protein level. After divergent selection for protein content, lines were evaluated for seed quality characters across three environments in Austria. The objectives of this research were to determine the roles of genetic background and the selection for protein content on food‐grade soybean traits. While seed protein content of adapted parents was between 395 and 420 g/kg, its range was from 410 to 490 g/kg for the high‐protein and from 390 to 450 g/kg for the food‐grade lines, respectively. However, food‐grade populations were superior in seed size and sucrose content and revealed different correlation patterns between quality traits as compared to high‐protein populations, which demonstrates their usefulness for developing soybeans with improved quality.     

Quantitative trait locus analysis for soybean (Glycine max) seed protein and oil concentrations using selected breeding populations

Soybean seed protein and oil concentrations are important traits that directly affect the quality of soyfoods. Many studies and breeding programmes have been conducted to find major quantitative trait loci (QTL) that regulate protein and oil concentrations and to develop soybean cultivars with high protein and/or oil content. The purpose of this study was to identify these QTL using a selected breeding population. The population was tested in field conditions over a period of 3 years. Seed protein and oil concentrations were measured each year. Single‐nucleotide polymorphisms (SNPs) were used to construct genetic map using a 180K SoyaSNP array, which identified 1,570 SNPs. We identified 12 QTL for seed protein, 11 for seed oil concentration and four for both traits. Among these, 17 QTL were closely mapped to previously reported QTL, whereas ten sites were novel. Several QTL were detected across at least two experimental years. These loci are good candidate QTL for optimal seed protein and oil concentrations. Our results demonstrate that favourable target QTL can be successfully identified using selected breeding populations.     

Genotypic basis of response to saliniity stress in some crosses of spring wheat Triticum aestivum L.

Wheat genotypes HD 2009, WH 157 and Kh 375 and their six F₁ crosses were evaluated for grain yield, biological yield and 1,000 grain weight under four levels of salinity (ECe 2.1, 6.2, 8.5 and 10.6 dS m^-1) in lysimeter type microplots. Parents Kh 375, WH 157 and HD 2009 were tolerant, moderately tolerant and sensitive to salinity, respectively. Reciprocal differences for salinity tolerance occurred for grain yield and 1,000 grain weight. The sensitive parent response was partially dominant whereas the salinity tolerant parent showed partial dominance for yield potential. Salinity tolerance and yield potential appeared to be controlled by different gene complexes. The cross Kh 375 × HD 2009 should provide transgressive segregants combining high yield potential with high salt tolerance. This revised version was published online in July 2006 with corrections to the Cover Date.     

冀豆12遗传背景导入系蛋白、脂肪含量分布特征

以高蛋白品种冀豆12为受体亲本,不同来源、不同蛋白脂肪含量的大豆种质资源为供体亲本,构建了28个组合BC2F1后代群体,分析冀豆12遗传背景导入系后代蛋白、脂肪含量分布特征。结果表明,28个后代群体均有蛋白含量超高亲个体,超高亲个体比例介于4.0%～68.2%之间,超高亲比例≥40%的组合有18个,占64.3%,BC2F1后代群体蛋白含量以超高亲和偏高亲类型组合为主。而脂肪含量分布特征恰相反,BC2F1后代群体脂肪含量以超低亲和偏低亲类型组合为主,超高亲个体比例介于0～67.4%,超高亲个体比例≥40%的组合有7个,占25.0%,9个组合无超高亲后代。表明以冀豆12为遗传背景通过有限回交易选育高蛋白含量品种,而不易选育高脂肪含量品种。本研究结果为利用冀豆12培育高蛋白品种提供了依据。