不同地区啤酒大麦品种农艺性状鉴定与分类研究

为筛选可用于云南啤酒大麦改良的优良种质材料,对107份不同来源啤酒大麦品种的农艺性状进行鉴定与分类研究.结果表明,云南啤酒大麦的主要特点是成熟期长,分蘖较强,植株高,旗叶面积大,穗长、穗粒数、穗粒重、千粒重、秸秆干重中等,穗草比较高;国外引进品种株型中等紧凑,叶片细长,穗粒重和千粒重较大,穗草比适中.从中筛选出19份性状优良的材料.通过聚类分析,可将供试材料划分成4个性状不同的类群,各类群的农艺性状差异明显,有利于育种目标材料的选择.     

The use of AFLPs to examine genetic relatedness in barley

The generation of AFLPs in spring barley cultivars provided genetic information relating to the development of the crop in the UK since 1953. Principal co-ordinate (PCO) analysis of genetic similarities (gs) confirmed the marked contrast in the cultivars used in the 1970s and 1980s. The earliest cultivars, many derived from Proctor, were succeeded by tall-strawed, disease-resistant types with high yield but poor malting potential. In the 1980s they were in turn replaced by short-strawed cultivars with excellent yield and good malting quality, which originated from Triumph. A PCO plot of gs provided insight into the effects of selection for disease resistance and the antagonism between malting quality and particular resistance genes. The analysis of gs was more useful than pedigrees and estimates of kinship in revealing the genetic relationship between cultivars. Theoretical considerations for maximising the efficiency of an AFLP genotyping programme are discussed in the context of the number of primer pairs required to distinguish genotypes at varying levels of similarity.     

Breeding progress,genotypic and environmental variation and correlation of quality traits in malting barley in German official variety trials between 1983 and 2015

Key message

Evaluation of breeding progress for spring barley varieties in Germany showed that both grain yield and malting quality were considerably improved during the last 33 years, and that genetic effects of protein concentration and malting traits were not associated.

Abstract

Based on historical data, this study aimed to investigate yield potential and malting quality of 187 varieties tested and released in German registration trials to evaluate the value for cultivation and use (VCU) during 1983–2015, and to quantify the environmental variability and the association among traits. We used mixed linear models with multiple linear regression terms to dissect genetic and non-genetic trend components. Grain yield increased by 43% (23.4 dt ha^?1) in VCU trials and 35% (14.0 dt ha^?1) on-farm relative to 1983. All yield components contributed significantly. Malting quality was also considerably improved by 2.3% for extract content up to 25.1% for friability, relative to 1983, nearly completely due to new varieties. Total variability of individual traits was very different between traits (2.4–24.4% relative to 1983). The relative influence of genotypes on total variation was low for grain yield and its components, whereas it was considerably larger for other traits. We found remarkable differences between phenotypic and genetic correlation coefficients for grain yield and protein concentration with malting traits. The observed positive phenotypic relation between grain yield and malting quality can be attributed to a shift of selection and environmental effects, but genetic correlations showed a negative association. Genetic effects of protein concentration and malting quality were not correlated indicating that both were not genetically linked. Considerable yield progress and improvement of malting quality were achieved despite of their weak to moderate negative genetic dependence.

     

Functional association between malting quality trait components and cDNA array based expression patterns in barley (Hordeum vulgare L.)

We developed an approach for relating differences in gene expression to the phenotypic variation of a trait of interest. This allows the identification of candidate genes for traits that display quantitative variation. To validate the principle, gene expression was monitored on a cDNA array with 1400 ESTs to identify genes involved in the variation of the complex trait malting quality in barley. RNA profiles were monitored during grain germination in a set of 10 barley genotypes that had been characterized for 6 quality-associated trait components. The selection of the candidate genes was achieved via a correlation of dissimilarity matrices that were based on (i) trait variation and (ii) gene expression data. As expected, a comparison based on the complete set of differentially-expressed genes did not reveal any correlation between the matrices, because not all genes that show differential expression between the 10 cultivars are responsible for the observed differences in malting quality. However, by iteratively taking out one gene (with replacement) and re-computing the correlation, those genes that are positively contributing to the correlation could be identified. Using this procedure between 17 and 30 candidate genes were identified for each of the six malting parameters analysed. In addition to genes of unknown function, the list of candidates contains well-known malting-related genes. Five out of eight mapped candidate genes display linkage to known QTLs for malting quality traits. The described functional association strategy may provide an efficient link between functional genomics and plant breeding.     

Genetic analysis of grain and malt quality in an elite barley population

Quantitative trait loci (QTLs) associated with grain weight, grain width, kernel hardness and malting quality were mapped in a doubled haploid population derived from two elite Australian malting barley varieties, Navigator and Admiral. A total of 30 QTLs for grain weight, grain width and kernel hardness were identified in three environments, and 63 QTLs were identified for ten malting quality traits in two environments. Three malting quality traits, namely β-amylase, diastatic power and apparent attenuation limit, were mainly controlled by a QTL linked to the Bmy1 gene at the distal end of chromosome 4H encoding a β-amylase enzyme. Six other malting quality traits, namely α-amylase, soluble protein, Kolbach index, free amino-acid nitrogen, wort β-glucan and viscosity, had coincident QTL clustered on chromosomes 1HS, 4HS, 7HS and 7HL, which demonstrated the interdependence of these traits. There was a strong association between these malt quality QTL clusters on chromosomes 1HS and 7HL and the major QTL for kernel hardness, suggesting that the use of this trait to enable early selection for malting quality in breeding programs would be feasible. In contrast, the majority of QTLs for hot-water extract were not coincident with those identified for other malt quality traits, which suggested differences in the mechanism controlling this trait. Novel QTLs have been identified for kernel hardness on chromosomes 2HL and 7HL, hot-water extract on 7HL and wort β-glucan on 6HL, and the resulting markers may be useful for marker-assisted selection in breeding programs.     

Genome-Wide Association Mapping for Kernel and Malting Quality Traits Using Historical European Barley Records

Malting quality is an important trait in breeding barley (Hordeum vulgare L.). It requires elaborate, expensive phenotyping, which involves micro-malting experiments. Although there is abundant historical information available for different cultivars in different years and trials, that historical information is not often used in genetic analyses. This study aimed to exploit historical records to assist in identifying genomic regions that affect malting and kernel quality traits in barley. This genome-wide association study utilized information on grain yield and 18 quality traits accumulated over 25 years on 174 European spring and winter barley cultivars combined with diversity array technology markers. Marker-trait associations were tested with a mixed linear model. This model took into account the genetic relatedness between cultivars based on principal components scores obtained from marker information. We detected 140 marker-trait associations. Some of these associations confirmed previously known quantitative trait loci for malting quality (on chromosomes 1H, 2H, and 5H). Other associations were reported for the first time in this study. The genetic correlations between traits are discussed in relation to the chromosomal regions associated with the different traits. This approach is expected to be particularly useful when designing strategies for multiple trait improvements.     

Dissection of a malting quality QTL region on chromosome 1 (7H) of barley

Malting and brewing are major uses of barley (Hordeum vulgare L.) worldwide, utilizing 30–40% of the crop each year. A set of complex traits determines the quality of malted barley and its subsequent use for beer. Molecular genetics technology has increased our understanding of genetic control of the many malting and brewing quality traits, most of which are quantitatively inherited. The objective of this study was to further dissect and evaluate a known major malting quality quantitative trait locus (QTL) region of about 28 cM on chromosome 1 (7H). Molecular marker-assisted backcrossing was used to develop 39 isolines originating from a Steptoe / Morex cross. Morex, a 6–row malting type, was the donor parent and Steptoe, a 6–row feed type, was the recurrent parent. The isolines and parents were grown in four environments, and the grain was micro-malted and analyzed for malting quality traits. The effect of each Morex chromosome segment in the QTL target region was determined by composite interval mapping (CIM) and confirmed and refined by multiple interval mapping (MIM). One QTL was resolved for malt extract content, and two QTLs each were resolved for -amylase activity, diastatic power, and malt -glucan content. One additional putative malt extract QTL was detected at the plus border of the target region by CIM, but not confirmed by MIM. All QTLs were resolved to intervals of 2.0 to 6.4 cM by CIM, and to intervals of 2.0 cM or less by MIM. These results should facilitate marker-assisted selection in breeding improved malting barley cultivars.     

QTL mapping reveals genetic architectures of malting quality between Australian and Canadian malting barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L.)

Australia and Canada are major exporters of malting barley (Hordeum vulgare L.), with Baudin from Australia and AC Metcalfe from Canada being the benchmark varieties for premium malting quality in the past 10 years. We used the barley doubled haploid population derived from a cross of Baudin and AC Metcalfe to map quantitative trait loci (QTLs) for malting quality. The results revealed different genetic architectures controlling malting quality for the two cultivars. Sixteen QTLs were identified and located on chromosomes 1H, 2H, 5H and 7H. The Australian barley Baudin mainly contributed to the malting quality QTL traits of high diastatic power and high β-glucanase on chromosome 1H, while Canadian barley AC Metcalfe mainly contributed to the QTL traits of high hot water extract, high free amino nitrogen, high α-amylase and low malt yield in chromosome 5HL telomere region. This study demonstrated the potential to breed new barley varieties with superior malting quality by integrating genes from Australian and Canadian malting barley varieties. This paper also provides methods to anchor traditional molecular markers without sequence information, such as amplified fragment length polymorphism markers, into the physical map of barley cv. ‘Morex’.     

The consequences, for malting quality, of Hordeum laevigatum as a source of mildew resistance in barley breeding

Barley cultivars which include Hordeum laevigatum in their pedigree, have harder endosperms, higher β-glucan and lower α-amylase levels than do other cultivars of poor malting quality. Good malting cultivars are characterised by high hot water extracts, soft endosperms, low β-glucan and high α-amylase levels. Introgressions of exotic genes for mildew resistance, into breeding populations, may be accompanied by factors adversely affecting malting quality.     

Haplotyping barley <Emphasis Type="Italic">bmy1</Emphasis> using the SNaPshot assay

The allelic status at bmy1, which encodes the enzyme β-amylase 1 in the barley grain, has an important influence over a cultivar’s malting quality. Changes in the malting process have been responsible for the need to improve the thermostability of this enzyme. We have compared a published bmy1 haplotyping assay based on TDI-FRET (template-directed dye-terminator incorporation fluorescence resonance energy transfer) with a SNaPshot protocol by jointly analysing a set of 21 cultivars of known haplotype. The two methods gave the same result, but the SNaPshot assay was easier to interpret. The SNaPshot assay was therefore used to haplotype the Czech malting barley core collection with respect to bmy1. The old Czech cultivar Kasticky was the only entry identified as carrying the high thermostability haplotype, with the remainder carrying either the intermediate or the low thermostability haplotypes. Older materials were the most variable in terms of bmy1 haplotype, but the majority carried the intermediate type. Most of the descendants of cv. Diamant carried the low thermostability haplotype. The most recently released cultivars recommended for the brewing of Czech beer tend to carry the intermediate allele.     

Markers polymorphic among malting barley ( Hordeum vulgare L.) cultivars of a narrow gene pool associated with key QTLs

Barley used for malting is a fine-tuned organism, and it requires breeding within narrow gene pools for realistic cultivar enhancement. Significant phenotypic advance within such narrow gene pools has been achieved and the necessary genetic variability for breeding progress has been documented, but it was not well understood. This study was conducted to further characterize detectable genetic variability present within a select set of four closely related malting barley cultivars using three types of molecular markers: RFLP, PCR-RAPD and AFLP. The markers that identified polymorphism among the select malting cultivars tended to link with each other and to map in chromosomal regions associated with quantitative trait loci (QTLs) for agronomic and malting quality traits that differed among the four cultivars. Although RFLPs identified the least amount of polymorphism, the differences detected by the RFLPs best fit the chronology of the cultivars. These results indicate that a large amount of the genetic variability necessary for cultivar improvement may have originally been present in the breeding gene pool, but does not rule out de novo variation. Study of the populations from crosses within this narrow germplasm is needed to further elucidate the basis of the phenotypic variability found among these select barley cultivars.     

Studies on the patterns of nitrogen uptake and translocation to the grain of winter barley intended for malting

In five experiments carried out between 1987 and 1989, crops of winter barley were grown under a range of agronomic treatments to study the way the treatments influenced the patterns of nitrogen uptake, dry matter production, translocation of assimilates and malting quality. Rate and timing of nitrogen fertiliser had large effects. Increased and late applications invariably increased nitrogen concentration in the grain by affecting the partitioning of nitrogen between the vegetative and reproductive parts of the plant. Foliar fungal diseases also affected nitrogen partitioning, apparently by interfering with the transport of stored nitrogen to the developing grain. Seasonal influences, especially rainfall during the grain filling period, were found to have profound effects on the pattern of change in ear and grain nitrogen concentration, and hence malting quality. The ability to predict final grain nitrogen concentration from measurements made earlier in the season is not therefore likely to be feasible.     

Molecular marker-assisted selection for enhanced yield in malting barley

Brewers are reluctant to change malting barley (Hordeum vulgare ssp. vulgare L.) cultivars due to concerns of altered flavor and brewing procedures. The U.S. Pacific Northwest is capable of producing high yielding, high quality malting barley but lacks adapted cultivars with desirable malting characteristics. Our goal was to develop high yielding near isogenic lines that maintain traditional malting quality characteristics by transferring quantitative trait loci (QTL) associated with yield, via molecular marker-assisted backcrossing, from the high yielding cv. Baronesse to the North American two-row malting barley industry standard cv. Harrington. For transfer, we targeted Baronesse chromosome 2HL and 3HL fragments presumed to contain QTL that affect yield. Analysis of genotype and yield data suggests that QTL reside at two regions, one on 2HL (ABG461C-MWG699) and one on 3HL (MWG571A-MWG961). Genotype and yield data indicate that additional Baronesse genome regions are probably involved, but need to be more precisely defined. Based on yield trials conducted over 22 environments and malting analyses from 6 environments, we selected one isogenic line (00-170) that has consistently produced yields equal to Baronesse while maintaining a Harrington-like malting quality profile. We conclude there is sufficient data to warrant experiments testing whether the 2HL and 3HL Baronesse QTL would be effective in increasing the yield of other low yielding barley cultivars.     

Genetic diversity among cultivars of spring barley revealed by random amplified polymorphic DNA (RAPD)

RAPD (random amplified polymorphic DNA) polymorphism was studied in 23 malting and non-malting spring barley cultivars included in the official list of Polish cultivated varieties. Twenty-four 10-mer primers were tested in each cultivar, giving altogether 149 amplification products, 45% of which were polymorphic. The number of polymorphic bands revealed by one primer ranged from 1 to 6, with an average of 2.8. Genetic distance for all pairs of compared varieties was estimated and a dendrogram was constructed using unweighted pair group method of arithmetic means. The genetic distance between cultivars ranged from 0.11 for cvs. Apex and Bryl to 0.62 for cvs. Orthega and Madonna. Of the seven malting cultivars only two (Brenda and Stratus) formed one group at D = 0.25. The genetic distance between cvs. Brenda and Scarlett, especially recommended for brewery, was equal to 0.34. The detected polymorphism appeared to be sufficient for assessing genetic distances between cultivars, but on the basis of this polymorphism groups of malting and non-malting cultivars were not clearly distinguished.     

Marker-assisted analysis of three grain yield QTL in barley (Hordeum vulgare L.) using near isogenic lines

Three previously identified grain yield quantitative trait loci (QTL) on chromosomes 2S(2HS), 3C(3HC) and 5L(1HL), designated QTL-2S, QTL-3 and QTL-5L, respectively, were evaluated for their potential to increase yields of high-quality malting barley without disturbing their favorable malting quality profile. QTL mapping of yield related traits was performed and near-isogenic lines (NILs) were developed. QTL for plant height, head shattering, seed weight and number of rachis nodes/spike were detected in the QTL-3 region. NILs developed by introgressing QTL-3 from the high-yielding cv. Steptoe to the superior malting quality, moderate-yielding cv. Morex acquired reduced height, lodging and head shattering features of Steptoe without major changes in malting quality. The yield of NILs, measured by minimizing the losses due to lodging and head shattering, did not exceed that of Morex. Steptoe NILs, with the Morex QTL-2S region, flowered 10 days later than Steptoe but the grain yield was not changed. None of the 3 QTL studied altered the measured yield of the recipient genotype, per se, although QTL 2S and QTL-3 affected yield-related traits. We conclude that these yield QTL must interact with other genes for full expression. Alternatively, they affect the harvestable yield through reduced lodging, head shattering, and/or altered flowering time.     

Development of DNA markers associated with beer foam stability for barley breeding

Traits conferring brewing quality are important objectives in malting barley breeding. Beer foam stability is one of the more difficult traits to evaluate due to the requirement for a relatively large amount of grain to be malted and then the experimental costs for subsequent brewing trials. Consequently, foam stability tends to be evaluated with only advanced lines in the final stages of the breeding process. To simplify the evaluation and selection for this trait, efficient DNA makers were developed in this study. Previous studies have suggested that the level of both of the foam-associated proteins Z4 and Z7 were possible factors that influenced beer foam stability. To confirm the relationship between levels of these proteins in beer and foam stability, 24 beer samples prepared from malt made from 10 barley cultivars, were examined. Regression analyses suggested that beer proteins Z4 and Z7 could be positive and negative markers for beer foam stability, respectively. To develop DNA markers associated with contents of proteins Z4 and Z7 in barley grain, nucleotide sequence polymorphisms in barley cultivars in the upstream region of the translation initiation codon, where the promoter region might be located were compared. As a result, 5 and 23 nucleotide sequence polymorphisms were detected in protein Z4 and protein Z7, respectively. By using these polymorphisms, cleaved amplified polymorphic sequence (CAPS) markers were developed. The CAPS markers for proteins Z4 and Z7 were applied to classify the barley grain content of 23 barley cultivars into two protein Z4 (pZ4-H and pZ4-L) and three protein Z7 (the pZ7-H, pZ7-L and pZ7-L2) haplotypes, respectively. Barley cultivars with pZ4-H showed significantly higher levels of protein Z4 in grain, and those with pZ7-L and pZ7-L2 showed significantly lower levels of protein Z7 in grain. Beer foam stability in the cultivars with pZ4-H and pZ7-L was significantly higher than that with pZ4-L and pZ7-H, respectively. Our results indicate that these CAPS markers provide an efficient selection tool for beer foam stability in barley breeding programs.     

Mildew-resistant mutants induced in North American two- and six-rowed malting barley cultivars

Mildew-resistant mutants were induced with sodium azide in three North American malting barley cultivars, two in the six-rowed Ursula (URS1 and URS2), one in the six-rowed Gertrud (GER1), and one in the two-rowed Prudentia (PRU1). Two of the mutants, URS1 and PRU1, showed complete resistance and were shown to have two new alleles at the mlo locus; these were designated, respectively, mlo31 and mlo32. Mutant URS2, showing partial resistance, was inherited as a dominant gene, but was not an allele at the Mla locus. The mean yield of each mutant was higher than that of its parental line, but yield levels varied across environments, although this was independent of the severity of the mildew attack. Other reasons, for example, the severity of the necrotic lesions in the mutants, may account for yield variations. The malting quality of the GER1 mutant proved similar to that of Gertrud, but both URS1 and URS2 showed lower malt extract than Ursula. This lower extract might be due to the smaller grain size of the mutants that could, in turn, result from necrotic lesions in the leaves, as implied by the effects on grain yield.Communicated by G. Wenzel     

强筋与弱筋小麦籽粒蛋白质组分与加工品质对灌浆期弱光的响应

选用强筋小麦品种济麦20和弱筋小麦品种山农1391,在大田试验条件下,分别于籽粒灌浆前期(花后6—9 d)、中期(花后16—19 d)和后期(花后26—29 d)对小麦进行弱光照处理,研究了籽粒产量、蛋白质组分及加工品质的变化。灌浆期弱光显著降低小麦籽粒产量,灌浆中期对济麦20和灌浆后期对山农1391的产量降幅最大。弱光处理后,籽粒氮素积累量及氮素收获指数减少。但弱光使籽粒蛋白质含量显著升高,其中灌浆中期弱光升幅最大,原因可能是由于其粒重降低造成的。弱光对可溶性谷蛋白无显著影响,但增加不溶性谷蛋白含量,使谷蛋白聚合指数显著升高,面团形成时间和稳定时间亦升高,籽粒灌浆中、后期弱光对上述指标的影响较前期大。灌浆期短暂的弱光照对改善强筋小麦粉质仪参数有利,但使弱筋小麦变劣;并均伴随籽粒产量的显著降低这一不利影响。     

Mapping and sequence analysis of barley hordoindolines

Barley (Hordeum vulgare L.) cultivars vary in traits such as grain hardness and malt quality. However, little is known about the genetic basis of these grain quality traits in barley, while more is known about the basis of grain hardness in wheat (Triticum aestivum L.). Puroindolines are endosperm-specific proteins found in wheat and barley, as well as other members of the Triticeae. In wheat, variation of puroindoline sequence is associated with most of the variability in wheat grain texture. However, no information exists on sequence variation of the barley homologs of puroindolines, the hordoindolines. We have therefore chosen to isolate and characterize the hordoindoline (hin) sequences of eight North American barley cultivars. The barley sequences contain numerous non-conservative amino-acid substitutions relative to their wheat counterparts. However, no significant rearrangements were found in either hinA or hinB of barley. Three hinA and two hinB sequence types were found among the eight barley cultivars examined, indicating substantial allelic variation at this locus. The hinB sequence variability was used to map hinB to the short arm of chromosome 5H in a Steptoe/Morex mapping population, which is coincident with the previously mapped location of hinA and Gsp (grain-softness protein). This chromosomal location also coincides with a small barley malt-extract QTL, suggesting that hordoindoline sequence variation may play a small role in barley grain quality. Efforts to correlate barley seed textural differences and malting quality with hordoindoline sequence type are ongoing. Received: 25 May 2000 / Accepted: 21 September 2000