Effect of environment and genotype on bread-making quality of spring-sown spring wheat cultivars in China

Improvement of end-use quality in bread wheat depends on a thorough understanding of current wheat quality and the influences of genotype (G), environment (E), and genotype by environment interaction (G × E) on quality traits. Thirty-nine spring-sown spring wheat (SSSW) cultivars and advanced lines from China were grown in four agro-ecological zones comprising seven locations during the 1998 and 1999 cropping seasons. Data on 12 major bread-making quality traits were used to investigate the effect of G, E, and G × E on these traits. Wide range variability for protein quantity and quality, starch quality parameters and milling quality in Chinese SSSW was observed. Genotype and environment were found to significantly influence all quality parameters as major effects. Kernel hardness, flour yield, Zeleny sedimentation value and mixograph properties were mainly influenced by the genetic variance components, while thousand kernel weight, test weight, and falling number were mostly influenced by the environmental variance components. Genotype, environment, and their interaction had important effects on test weight, mixing development time and RVA parameters. Cultivars originating from Zone VI (northeast) generally expressed high kernel hardness, good starch quality, but poor milling and medium to weak mixograph performance; those from Zone VII (north) medium to good gluten and starch quality, but low milling quality; those from Zone VIII (central northwest) medium milling and starch quality, and medium to strong mixograph performance; those from Zone IX (western/southwestern Qinghai-Tibetan Plateau) medium milling quality, but poor gluten strength and starch parameters; and those from Zone X (northwest) high milling quality, strong mixograph properties, but low protein content. Samples from Harbin are characterized by good gluten and starch quality, but medium to poor milling quality; those from Hongxinglong by strong mixograph properties, medium to high milling quality, but medium to poor starch quality and medium to low protein content; those from Hohhot by good gluten but poor milling quality; those from Linhe by weak gluten quality, medium to poor milling quality; those from Lanzhou by poor bread-making and starch quality; those from Yongning by acceptable bread-making and starch quality and good milling quality; and those from Urumqi by good milling quality, medium gluten quality and good starch pasting parameters. Our findings suggest that Chinese SSSW quality could be greatly enhanced through genetic improvement for targeted well-characterized production environments.     

Additive and epistatic effects of allelic variation at the high molecular weight glutenin subunit loci in determining the bread-making quality of breeding lines of wheat

Summary The relation has been studied between the high molecular weight glutenin (HMWg) subunit alleles and the bread-making quality of 226 lines of winter wheat (T. aestivum L.), grown in The Netherlands. The lines represented a wide range of genetic backgrounds, and had not been selected for quality, in contrast to the established varieties used by other authors.The variation in HMWg subunit genotypes accounted for about 20% of the total variation in loaf volume among the lines. Most important was the allelic variation at the Glu-D1 locus. The Glu-D1 allele encoding the subunits 5+10 was superior to its allelic counterpart, encoding 2+12. The difference in average of loaf volume between groups of lines containing 5+10 or 2+12 was negatively related with protein content of the flours. When protein content was below 9.2%, no effect of allelic variation at the Glu-D1 locus was present. Epistatic effects between the Glu-I loci also contributed to the variation in loaf volume of the lines: i.e. the effect of allelic variation at Glu-A1 and Glu-B1 depended on the allele present at the Glu-D1. The contribution of the epistatic effects was about half the contribution of the additive effects, and should therefore be included in predictive models for bread-making quality.     

Effects of homoeologous group 1 and 6 chromosomes of the Cappelle-Desprez (Bezostaya 1) substitution lines on aspects of bread-making quality of wheat

Summary The group 1 and 6 inter-varietal chromosome substitution lines of Cappelle-Desprez (Bezostaya 1) were intercrossed along with the donor and recipient varieties, Cappelle-Desprez and Bezostaya 1, to give 36 genetically different families. The analysis of the means of these families showed that variation in SDS-sedimentation volume fitted a predominantly additive model. There were no significant within or between chromosome interactions among the group 1 and 6 chromosomes. Nor was there any evidence for interactions between these chromosomes and those of the background. Significant dominance/within chromosome interactions amongst the background chromosomes were however detected. Some of the positive effects on SDS-sedimentation were associated with increased grain hardness. Chromosome effects on % grain protein were not correlated with SDS-sedimentation.     

Relationship between common wheat (Triticum aestivum L.) gluten proteins and dough rheological properties

This paper reports the correlation between the rheological properties of bread wheat dough and the types and quantities of endosperm proteins in 28 common wheat cultivars. Different methods were used to analyse the allelic composition of these cultivars and the relative quantities of the different proteins contributing to the gluten structure. Neither dough strength (W) nor tenacity/extensibility (P/L) correlated with allelic composition. Different wheats with the same allelic composition (i.e., with respect to glutenins) showed different rheological properties. The glutenins were the most influential components of W and P/L, especially the high molecular weight (HMW) glutenin subunits and in particular the type x form. These proteins seem to increase W and are the main constituents of the gluten network. The gliadins and low molecular weight (LMW) glutenin subunits appear to act as a “solvent”, and thus modify the rheological properties of the dough by either interfering with the polymerisation of the HMW glutenin subunits, or by altering the relative amounts of the different types of glutenin available. Thus, the protein subunits coded for by the alleles Glu-B1x7 and Glu-D1x5 stabilised the gluten network, whereas those coded for by Glu-B1x17 and Glu-D1x2 had the opposite effect. Dough properties therefore appear to depend on the glutenin/gliadins balance, and on the ratio of the type x and type y HMW proteins. The influence of external factors seems to depend on the allelic composition of each cultivar.     

Kernel hardness and baking quality of wheat — A genetic analysis using chromosome substitution lines

Summary An attempt was made to identify the chromosomal location of genetic control of a few components of wheat quality, using chromosome substitution lines of Cappelle Desprez, Cheyenne, Hope, and Timstein into the recipient variety Chinese Spring.Major factors for kernel hardness and increased baking absorption were found on chromosomes 5D of Cheyenne and Hope, and on 3B, 5D and 7D of Timstein. In Timstein, the presence of one of these chromosomes sufficed to make the wheat kernels hard.Factors for favourable dough properties were identified on a few other chromosomes, different in various varieties. These were 1A of Cappelle Desprez and Cheyenne, 3B of Hope, and 2D of Timstein. All but one of these chromosomes showed an increase in loaf volume to a level in-between those of the recipient variety Chinese Spring and the donor varieties. No relationship was found between kernel hardness and dough-making and baking properties.It was assumed that wheat quality is due to a combination of kernel hardness and favourable dough-making properties. As the genes for these factors are located on different chromosomes, it should not be too difficult to introduce both factors in existing varieties with poor baking properties. In a wheat breeding programme, the quality of new lines can be assessed in a rather simple way by determining kernel hardness and dough stability.     

Selection for yield in small plots of spring wheat

Summary Four selection experiments were sown: single-row plots with single row-spacing (20 cm), single-row plots with double row spacing (40 cm), three-row plots and six-row plots both with single row-spacing (20 cm). Selection entries were mimiced by 16 different varieties or advanced breeding lines, which were also sown in a yield trial. Each experiment was laid out as a 4-times replicated randomized block design. Row length was 2 m. Alley borders and border-rows of multiple-row plots were harvested separately to evaluate the effects of different harvesting procedures on the selection efficiency. Removal of alley borders was found to be disadvantageous, since the gain in precision was more than offset by the loss in sampled area. Wide spacing of single-row plots improved the selection efficiency in comparison with normal spaced single-row plots. In multiple-row plots the selection efficiency was not improved by harvesting only the central rows.For gross plot yield (= yield of net plot + yield of alley borders) the differences in selection efficiency between the various selection plot-types were explained on the basis of the genetic variance, the environmental variance and the coefficient of genetic correlation with farm yield as determined in the yield trial.     

The effect of indirect tests for grain quality on the grain yield and industrial quality of bread wheat

The objective of this study was to examine the effect of selection using three indirect tests for grain quality on grain yield and dough, and baking properties, measured as alveograph strength, alveograph tenacity/extensibility ratio and loaf volume. The three tests were flour protein content, flour sedimentation and high molecular weight glutenin subunits. Of the indirect tests used for grain quality, sodium dodecyl sulphate (SDS)‐sedimentation allowed the highest intensity of selection for a combined trait index of the target grain quality and grain yield characteristics. The top 48% of the material could be retained on the basis of SDS‐sedimentation, resulting in retention of atleast two‐thirds of the top 10% of genotypes for the combined trait index. Flour protein percentage, a weighted high molecular weight glutenin index and an index combining all the indirect tests—flour protein, SDS‐sedimentation and high molecular weight glutenins—gave selection intensities of 61%, 64% and 55%, respectively, for the combined trait index. If the objective of selection is dough strength alone, then a weighted index of all indirect traits (flour protein, SDS‐sedimentation and high molecular weight glutenins) provided the highest selection intensity (26%). Other selection intensities for individual target traits were 24% for the prediction of loaf volume from flour protein, 40% for the prediction of tenacity/ extensibility ratio using SDS‐sedimentation and 68% for the prediction of grain yield using SDS‐sedimentation.     

An evaluation of seed and seedling drought tolerance screening tests in wheat

Summary A series of experiments was performed in order to evaluate the significance of seed germination and seedling growth in osmotic media as screening methods for drought tolerance.Ten spring wheat (Triticum aestivum L. em Thell.) and one durum wheat (Triticum durum Desf.) were tested under controlled environments, using polyethylene glycol-6000 (PEG) solutions as the moisture stress inducing media.Tolerance in the rate of endosperm utilization, under stress, prior to the onset of germination varied among cultivars.Germination rate or injury to germination at various concentrations of PEG differed significantly among cultivars. Cultivar rating with respect to injury to germination changed with stress levels. Injury to germination did not correlate with endosperm utilization rate in PEG or in water.Germinating seedlings were tolerant to extreme desiccation up to the stage of emergence of the first leaf from the coleoptile.Growth of photosynthesizing seedlings was monitored as they were carried through an increasing concentration gradient of PEG solutions, ranging from –5.9 to –11.3 bars of water potential. Cultivars significantly differed in seedling growth tolerance to increasing levels of water stress. Seedling growth tolerance across cultivars was not correlated with their germination responses under srress.It is concluded that tolerance to water stress in growing seedlings can be screened for by using PEG-containing nutrient solutions. It can not be predicted from germination tests in osmotica.Work was done under a US-Israel Binational Science Foundation (BSF) Grant no. 1654/78.Contribution from Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel, No. 192-E, 1979 series.     

Breeding for resistance to cereal cyst nematode in wheat II. Use of test tube cultures in selection

Summary The use of soil. naturally infested with Heterodera avenae, to select resistant heterozygotes in backcross progenies of wheat, was tested for reliability. Selfed progenies from plants selected as resistant were cultured monoxenically in test tubes with nematodes hatched from single cysts, while backcross progenies from the same parent plants were grown in pots of naturally infested soil. Cyst counts were made after two months' growth. The results showed that over 50% of the backcross lines, screened in previous generations with naturally infested soil, had been erroneously selected as resistant. The test tube cultures clearly differentiated lines carrying resistance from those which were susceptible and corroborated results from pot tests.     

Selection for biomass yield in wheat

Summary Biomass (above ground plant parts) yield may be a useful selection trait for yield improvement in wheat (Triticum aestivum L.). This study was conducted to estimate realized heritability of biomass yield and to determine the response to selection for high and low biomass yield in 8 genetically diverse populations of spring wheat under two production systems. Selections were made among the F₃ lines. Progenies of the selected lines were evaluated in replicated field tests in the F₄ generation under high fertility and low fertility production systems at Rampur, Nepal, in 1991. Fertility level had a significant effect on biomass yield, grain yield, effective tiller number, number of kernels per spike, thousand kernel weight, and harvest index. Selection in the F₃ for high and low biomass yield was effective in identifying F₄ lines with high and low biomass yield, respectively. Biomass yield differences between high and low selection groups in the F₄ generation, expressed as percent of the mean of the low selection group and averaged over the eight populations, were 53.9 and 36.5% higher than the mean of the low selection group under the high and the low fertility production systems, respectively. The corresponding figures for grain yield were 48.8 and 34.9% under the high and the low production systems, respectively. Also, selection for high biomass yield resulted in higher effective tiller number, and number of kernels per spike, but lower harvest index. Realized heritability estimates for biomass yield were greater at high fertility (range 0.49 to 0.85) than at low fertility (range 0.22 to 0.44). Biomass yield showed positive genotypic correlations with grain yield, effective tiller number, and number of kernels per spike but a negative correlation with harvest index. The results indicated that selection for high biomass yield should bring about positive improvements in biomass yield, grain yield, effective tiller number, and number of kernels per spike. The correlation between F₃ and F₄ generations suggested that biomass yield in the F₃ generation was a good predictor of biomass yield and grain yield in the F₄ generation. Selection for biomass yield in wheat should be made under the standard production system to obtain a realistic response.     

Honeycomb selection for yield in early generations of spring wheat

Summary The effectiveness of the honeycomb selection method for yield in spring wheat (Triticum aestivum L.) was evaluated using progenies from two wheat crosses, Glenlea x NB131 and Glenlea x Era. Honeycomb selection was carried out in the F₂ and F₃ generations, grown at the University of Manitoba in the summers of 1980 and 1981, respectively. In both generations, divergent selection was made for both high and low yield. Plants selected in the F₃ generation were entered in an F₄ yield test in the summer of 1982. Results of the experiment showed that honeycomb selection for yield in the F₂ and F₃ generations was effective in identifying parents of high- and low-yielding lines. F₃ plants from highyielding F₂ selections gave higher yields than those from low-yielding F₂ selections by 11.5% and 13.0% for Glenlea x NB131 and Glenlea x Era crosses, respectively. The F₄ yield test showed that high yielding selections from both crosses significantly outyielded by 8.9% low yielding selections and by 14.4% the unselected composite lines. It is concluded that the honeycomb selection method can be used for early generation selection in spring wheat.     

Polyacrylamide gel electrophoresis of wheat gliadins: the use of a moving boundary for improved resolution

Summary The process of the separation of gliadin proteins of wheat, using polyacrylamide gel electrophoresis, was studied in detail. It was shown that electrophoresis is a dynamic process during which the pH of the gel changes together with the potassium ion concentration. The most positive effect on the separation of the gliadins was generated by a moving front, which is a boundary between regions with a low concentration of K⁺ ions and a low pH, and region with a high concentration of K⁺ ions and a high pH, after optimization of the concentrations of the cations and anions in the electrode solutions. The finding was exploited for the development of an extremely simple electrophoresis system, in which buffers were not needed for obtaining a high resolution. The system was further improved by applying a stacking gel. The advantages of this system are discussed. This new approach may be useful for improving electrophoresis systems for other applications.     

Deficiency of individual high molecular weight glutenin subunits affords flexibility in breeding strategies for bread-making quality in wheat Triticum aestivum L.

High-molecular-weight (HMW) glutenin subunits in wheat Triticumaestivum L., allelic variation for which affects bread-making quality, areencoded by Glu-1 homoeoloci located on the homoeologous group1 chromosomes. Many alleles at Glu-B1 and Glu-D1 producetwo subunits, an x-type of low electrophoretic mobility in polyacrylamidegels, and a y-type of high mobility. In the current study, a combination ofnear isogenic lines of cultivar `Sicco' has been used to characterise theeffect upon quality of the absence of individual subunits 7 (Glu-B1x-type), 12 (Glu-D1 y-type) and, assuming an additive model ofsubunit action, 2 (Glu-D1 x-type). Absence of subunit 7 gave amoderate reduction in SDS-sedimentation volume, indicating its associationwith lower gluten strength (confirmed by Farinogram and Extensogramstudies), yet, from a full mixing input bake, a moderate increase in loafvolume and a considerable improvement in loaf score (an overall evaluationof loaf quality). Absence of subunit 12 gave a slightly larger reduction inSDS-volume, yet no change in loaf volume or score. Absence of bothsubunits 2+12 gave a larger reduction again in SDS-volume, a moderatereduction in loaf volume and a large reduction in loaf score. Absence ofsubunit 2 alone is therefore predicted to reduce SDS-volume, loaf volumeand score such that loss of this x-type subunit would lead to larger changesin quality parameters than loss of y-type subunit 12. A general conclusionof the study is that, while deficiency for HMW glutenin subunits generallyleads to reduced gluten strength and viscoelasticity, the resultantintermediate gluten strength may on occasions lead to improvements in loafperformance in situations where the base gluten strength is high. Theremay, then, be contexts in breeding programmes where selection fordeficiency would be a possible strategy for improving bread-making quality,adding to the flexibility available to the breeder. Somewhat unexpectedly,additional analysis found that, in the genetic background of cultivar `Sicco'used in this study, subunits 7+8 at Glu-B1 were indistinguishablefrom their allelic counterparts subunits 7+9 for virtually all characters, andthat subunits 2+12 at Glu-D1, while inferior in performance formixing properties compared to subunits 5+10, were associated with goodloaf characteristics.     

Agronomic and quality characteristics of spring wheat lines selected for protein content and protein yield

Summary High protein cultivars of spring wheat (Triticum aestivum L.) from eight foreign countries and the United States were used in crosses to provide progeny for a recurrent selection program. After two cycles of selection, 40 lines selected for protein yield and 16 lines selected fro protein percentage were evaluated with parents in yield nurseries at Bozeman, Montana.Lines selected for protein pereentage had the highest protein percentages, protein yields similar to the parents, and grain yields lower than the parents. Lines selected for protein yield had protein percentages intermediate between those of the parents and lines selected for protein percentage, but they had the highest protein yields and grain yields. Protein percentage and grain yield were negatively correlated and protein yield and grain yield were positively correlated for both groups of lines. The data tend to suggest that protein yield may a better selection criterion than protein percentage for plant breeders to use in improving protein productivity, although additional testing of this hypothesis is proposed.Milling and baking data showed transgressive improvement over the mean of the patents in many important quality aspects, indicating that good quality lines can be obtained from crosses involving poor to mediocre quality cultivars.Joint contribution of ARS, USDA and the Mont. Agric. Exp. Stn. Published with approval of the Director of the Mont. Agric. Exp. Stn. as Paper No. 1181, Journal Series. Received 19 June 1981.     

A test for linkage between Sr6 and Ra in wheat

Summary Previous evidence suggested that the gene for red auricle (Ra) might be linked to the gene Sr6 for stem rust resistance on chromosome 2D. However, tests indicated that Ra is not linked to either Sr6 or C, which is also on 2D.     

The effect ofGlu-B1 high molecular weight glutenin subunits on biscuit-making quality of wheat

Summary The aim of this study was to assess the effect of specificGlu-B1 HMW-GS on biscuit-making quality. Three soft spring wheat cultivars with the sameGlu-A1 andGlu-D1 HMW-GS, but differentGlu-B1 HMW-GS were used in crosses. F₂₄ derived lines were developed from these crosses.Glu-B1 HMW-GS 6+8 and 17+18; and 7+9 and 17+18 were compared. Lines with HMW-GS 6+8 versus those with HMW-GS 17+18 had a higher flour protein- and alveograph P/L ratio, shorter mixograph mixing time, more vitreous kernels, and a lower alveograph distensibility and strength (all values significant at p=0.05). Lines with HMW-GS 7+9 compared to those with 17+18 showed significant differences for flour extraction and biscuit diameter. The presence of HMW-GS 17+18 was significantly correlated with several biscuit-making quality characteristics in the Dirkwin/Zaragosa F₂₄ lines but not in the Waverley/Zaragosa F₂₄ lines, therefore the effect of HMW-GS 17+18 was modified by the genetic background in which they were expressed.     

Positive effect of the high-molecular-weight glutenin allele, Glu-D1d, on the bread-making quality of common wheat

The seed storage proteins of wheat flour are the determinants of bread‐making quality. Many cultivars having good bread‐making quality carry the Glu‐D1d allele responsible for the development of glutenin, a major seed storage protein. The Glu‐D1d allele was introduced into four leading Japanese wheat cultivars by recurrent backcrossing and the quality of these near‐isogenic lines (NILs) was evaluated by the sodium dodecyl sulphate sedimentation value of their flour. The values for the NILs were significantly higher than for the corresponding recipient cultivars. However, the values did not reach the level of the cultivar that had been used as the donor of the Glu‐D1d allele.     

Dough properties and bread-making quality-related characteristics of Yumechikara near-isogenic wheat lines carrying different Glu-B3 alleles

We investigated the relationships of three allelic variations in Glu-B3 (ab, g, and h) with dough properties and bread-making quality-related characteristics using near-isogenic lines (NILs) of ‘Yumechikara’ that commonly carry Glu-A1a, Glu-B1b, Glu-D1d, Glu-A3f, Glu-B3ab and Glu-D3a. Measurement of peak time (PT) in a 2-g mixograph indicated that Glu-B3g was the most effective for a strong dough property, followed by Glu-B3ab, with Glu-B3h being the least effective. The results of measurement of mixing time during bread-making were similar to those for PTs, i.e., the lines carrying Glu-B3g showed the longest mixing time, followed by those of Glu-B3ab, and those of Glu-B3h showed the shortest mixing time. Since two parameters of bread-making quality, loaf volume (LV) and specific loaf volume (SLV), were affected by flour protein contents in all groups of the Glu-B3 genotype, we compared the effects of the three Glu-B3 alleles on those parameters using analysis of covariance (ANCOVA) to remove the effect of protein content. The results indicated that the Glu-B3h group showed the largest SLV, followed by the Glu-B3ab group, and the Glu-B3g group showed the smallest SLV. These results suggest that the introduction of Glu-B3h into ‘Yumechikara’ makes it possible to breed varieties with good bread-making quality-related characteristics.     

A test of supposed mutants for stem rust resistance in wheat

Summary Eight stem rust (Puccinia graminis tritici Eriks. and Henn.) resistant lines (designated TICENA lines) that had been selected by Veiga et al. (1981) following gamma radiation of BH-1146 wheat (Triticum aestivum L.) were studied. Six of the lines were resistant to race 15B-1 of stem rust and susceptible to race 56, and proved to carry the gene Sr7a. TICENA 4 carries two unidentified genes, each giving resistance to one of the two races. TICENA 10 carries Sr6, Sr7a and an unidentified gene giving resistance to race 56 but not 15B-1. The results raise doubts about the supposed origin of the lines as mutants.     

A comparison of yield and yield component selection in winter wheat

Summary Three populations of winter wheat were formed by crossing Avrora to Sage, TAM W-101, and Danne. Approximately 10% of the F₂ plants from these crosses were selected for high and low levels of number of tillers per plant, number of kernels per spike, 1000-kernel weight, and grain yield. Forty-eight solid seeded F₃ lines obtained from the selected F₂ plants were then selected for high and low expressions of yield components and grain yield. Realized heritabilities were estimated. Indirect responses of yield to yield component selection and direct response to selection for grain yield were measured. Heritabilities were low for tiller number, number of kernels per spike and kernel weights but were high or intermediate for grain yield when selection occurred in the F₂ generation. When selection was practiced in the F₃ generation, heritabilities for tiller number and yield were low, but were intermediate to high for number of kernels per spike and kernel weight and high heritabilities were found for kernel weight. Selection for kernel weight often increased grain yield; however, direct selection for grain yields was usually as effective.Journal article no. J-4488 of the Oklahoma Agri. Exp. Stn., Stillwater, Oklahoma 74074.