白皮小麦收获前穗发芽及品种抗性机制探讨

对４８个白皮小麦种质及２个红皮小麦种质连续３年测定结果,品种间收获前穗发芽率,籽粒发芽率及收获后种子休眠特性差异显著。穗发芽敏感性因年份和种子发育时期而异,开花后３５－４０天较敏感,品种间差异大。休眠期长短与穗发芽率间呈极显著负相关。收获后的种子α－淀粉酶活性,降落值,与穗发芽率间分别呈极显著正相关和极显著负相关。     

Dormancy breaking procedures and the breeding of white-grained wheat with resistance to pre-harvest sprouting

Summary Gibberellic acid, cold (4°C) and a combination of these two treatments were tested for use in breaking dormancy in 27 lines of white-grained wheat with varying levels of resistance to pre-harvest sprouting. Germination increased in all lines treated with gibberellic acid. Dormancy could be broken with 1 M gibberellic acid. Response to cold varied. A combination of gibberellic acid and cold treatment was the most effective. This technique has been found useful in treating seed in a breeding program aimed at producing wheats with resistance to pre-harvest sprouting.     

Relationship between sprouting in wheat and embryo response to endogenous inhibition

Summary Seed dormancy in wheat (Triticum aestivum L. em Thell) is important for minimizing pre-harvest sprouting. To facilitate breeding cultivars that tolerate pre-harvest sprouting conditions, we assessed mode and magnitude of variation of seed dormancy among genotypes and investigated involvement of endogenous water-soluble inhibitor(s) in seed germination. Embryo bio-assays established that water-soluble inhibitor was ubiquitous among the wheat cultivars studied and did not diminish in quantity during after-ripening. Germination response of embryos was decreased by endogenous inhibitor, but the effect markedly declined as embryos aged at room temperature. Variation in dormancy among cultivars was primarily caused by differential response of their embryos to endogenous inhibitor. Gibberellic acid counteracted the initial inhibitory effect of endogenous inhibitor on germination but not the subsequent inhibitory effect on seedling growth. We concluded that pre-harvest sprouting resistance involves multiple factors, particularly embryo receptivity to endogenous inhibitor, and that variation in inhibitor quantity is not solely responsible for genotypic differences in susceptibility to pre-harvest sprouting. The possibility of additional approaches to breeding for pre-harvest sprouting resistance is indicated.Contribution no. 81-389-j, Department of Agronomy, Kansas State University, Manhattan, KS 66506, U.S.A.     

Breeding of white-grained wheats for Japan

The possibility of breeding white-grained wheats tolerant to pre-harvest sprouting under Japanese humid weather conditions is discussed. New genetic dormancy sources, such as, AUS1408, 8019R1 and RyuuMai7, were evaluated for seed dormancy in different weather conditions. Some white-grained dormant wheats showed a strong dormancy similar to that of red-grained dormant wheats in a greenhouse trial, in the field their dormancy expression was much less than the red wheats. Three populations involving crosses with these new sources for winter wheat breeding were examined under repeated selection for seed dormancy. Some dormant white-grained lines, as judged under glasshouse conditions, were developed. Again the level of dormancy in these lines was not sufficient compared to red dormant varieties in field trial. In order to develop truly superior dormant white-grained materials, one population involving crosses with two dormant varieties, AUS1408 and 8019R1, was examined under repeated selection. From these materials we succeeded in breeding lines that had not only a good dormancy but also showed adaptation to Japanese weather conditions, including earliness, scab resistance and good seed appearance. However again the dormancy of these lines in field trial was not sufficient compared to that of red wheats and there was not a clear difference for seed dormancy between breeding lines and their parents. We conclude therefore that more work involving the use of new genetic sources or new breeding techniques, will be necessary for breeding advanced lines that maintain a sufficient tolerance to PHS in humid Japanese weather condition. This revised version was published online in August 2006 with corrections to the Cover Date.     

小麦籽粒休眠Vp1-B1基因的等位变异检测与分离

Vp1基因是控制籽粒休眠性的重要基因之一,检测该基因的等位变异类型,可以进一步理解籽粒休眠以及穗发芽抗性的遗传控制机理.本文根据GenBank中小麦Vp1-B1基因序列设计特异引物检测我国小麦微核心种质以及地方品种和推广品种,结果表明,本研究除了发现已报道的3种等位变异类型,分别为Vp1-B1a、Vp1-B1b和Vp1-B1c,另外检测到一种新的变异类型,暂命名为Vp1-B1x.经改良的变性PAGE凝胶电泳检测以及序列比对分析表明,Vp1-B1x与Vp1-B1c基因的序列相似性最高,达99%;其在第3内含子区域发生"ATAT"4个碱基的插入以及4个SNP,但是该等位类型在所检测的品种中分布较少,其与籽粒休眠水平及穗发芽抗性之间的关系尚待进一步验证.     

Mapping quantitative trait loci controlling pre-harvest sprouting resistance in a red × white seeded spring wheat cross

Hard white wheat (Triticum aestivum L.) is a value-added product because of its processing advantages over red wheat; however, white wheat tends to be more susceptible to pre-harvest sprouting (PHS). To identify quantitative trait loci (QTLs) associated with PHS tolerance, we developed a doubled haploid (DH) mapping population from the cross AC Domain (red seeded) × White-RL4137 (white seeded). A genetic map was constructed using microsatellite markers located on chromosome groups 3, 4, 5 and 6. A population of 174 DH lines was characterized for important aspects of PHS including sprouting index, germination index, Hagberg falling number and seed coat colour. A total of 11 QTLs were identified on group 3 chromosomes and on chromosome 5D. Seven QTLs associated with the PHS traits were found to be co-incident with seed coat colour on chromosomes 3A, 3B and 3D. The 5D PHS QTL was notable because it is independent of seed coat colour.     

Genetic analysis of pre-harvest sprouting in a durum wheat cross

Pre-harvest sprouting of durum wheat (Triticum turgidum L. var durum) reduces commercial grade, although the actual effects on processing quality are controversial. Little is known about the genetics of the dormancy component of pre-harvest sprouting resistance in durum. We studied the segregation of dormancy in 98 recombinant inbred lines from a cross of a relatively non-dormant line, CI13102, with a moderately dormant line, Kyle. The lines and parents were grown in field tests over three years, 1996, 1997 and 1998. Spikes were collected at approximately 20% moisture and stored at −23 ∘C. Hand-threshed grain of the lines was germinated, and number of seeds germinated was counted each day. A germination resistance index was calculated to characterize dormancy. Dormancy appeared to be complexly inherited in this cross. Lines were observed that were significantly (P < 0.05) more dormant than the parents. The lines transgressive for dormancy expressed in different combinations of the three environments, indicating an environmental interaction. DNA of lines and parents was tested with simple sequence repeat primers and AFLPs that were used in quantitative trait loci (QTL) analysis of dormancy. Significant QTLs for dormancy were found, with the most notable being on chromosome 1A, where other QTLs for pre-harvest sprouting resistance have been reported in common wheat.     

Assessment of genetic diversity in synthetic hexaploid wheats and their Triticum dicoccum and Aegilops tauschii parents using AFLPs and agronomic traits

Synthetic hexaploid wheats are of interest to wheat breeding programs, especially for introducing new genes that confer resistance to biotic and abiotic stresses. A group of 54 synthetic hexaploid wheats derived from crosses between emmer wheat(Triticum dicoccum, source of the A and B genomes) and goat grass (Aegilops tauschii, D genome donor) were investigated for genetic diversity. Using the AFLP technique, dendrograms revealed clear grouping according to geographical origin for the T. dicoccum parents but no clear groups for the Ae. tauschii parents. The geographical clustering of the T. dicoccum parents was also reflected in the dendrogram of their derived synthetic hexaploids. Diversity of the T. dicoccum parents and their derived synthetic hexaploids was further evaluated by measuring 18morphological and agronomic traits on the plants. Clustering based on morphological and agronomic data also reflected geographical origin. However, comparison of genetic distances obtained from AFLP and agronomic data showed no correlation between the two diversity measurements. Nevertheless, similarities among major clusters with the two systems could be identified. Based on percentage of polymorphic markers, the synthetic hexaploids had a considerably higher level of AFLP diversity (39%) than normally observed in cultivated hexaploid wheat (12–21%). This suggests that synthetic hexaploid wheats can be used to introduce new genetic diversity into the bread wheat gene pool. This revised version was published online in July 2006 with corrections to the Cover Date.     

四川省小麦品种(系)穗发芽抗性评价

穗发芽严重影响小麦产量和品质,受到生长环境、种皮颜色和遗传基因的共同控制。由于四川地区在小麦收获季节容易出现高温高湿的气候条件,是小麦穗发芽的常发和重发区。依据材料的穗发芽抗性,有目的性地配制杂交组合选育抗性品种是控制小麦穗发芽的重要策略,而系统鉴定分析各优良品种及品系的穗发芽抗性是亲本选配的重要依据。本研究调查分析了242份2000年来四川省审定小麦品种及即将成为未来几年新品种的2017—2018 年四川省区试品系的田间穗发芽情况,发现分别有23(9.5%)和67 (27.7%)份材料具有强（穗发芽率低于5%）和高（穗发芽率介于5%~15%）的穗发芽抗性。这为穗发芽抗性育种亲本的选择提供了参考;并且发现穗发芽抗性在不同类型材料中存在差异：2000 年来审定品种＜区试普通小麦品系＜区试早播早熟小麦品系＜区试特殊用途小麦品系,穗发芽抗性在未来几年品种中将有所增强;种皮颜色对穗发芽抗性具有强烈影响,随着种皮颜色加深,穗发芽抗性越强,但仍发现3 份（‘川育26’、‘川辐9 号’、‘川辐17 号’）白皮小麦材料具有强的穗发芽抗性,为白皮小麦抗穗发芽育种提供参考。其次发现在四川环境下,早播可能会提高小麦品种的穗发芽抗性。     

Crossability of tetraploid and hexaploid wheats with ryes for primary triticale production

Summary Crossability of wheat and rye was investigated during thirteen crop cycles in two contrasting locations to 1) evaluate tetraploid and hexaploid wheat parents in crosses with rye, 2) identify genotypes with high crossability and 3) assess the impact of environment on seed development. The majority of the tetraploid wheats crossed with rye had seed set around 20%, but very low embryo viability. Several wheat genotypes with seed set above 50% were identified. The hexaploid wheats crossed with rye showed poor seed set, but plant recovery was relatively high. The majority of the hexaploid wheats with highest seed set (20–30%) were from China. The results suggest differences in crossability between the rye populations, and wheat species by rye interactions. The crossability of the tetraploid and hexaploid wheats was affected by climate in the two locations.     

Cereal comparative genetics and preharvest sprouting

Most genes in hexaploid bread wheat are triplicated. Knowledge of the relationships between the three genomes then allows us to build consensus maps of loci controlling any trait. In this paper we show such a map of some of the major genes and QTL effects that have been reported to be associated with pre-harvest sprouting. The result highlights regions of the genome that have featured in several studies and possible links between QTL and major genes. The same analysis can be extended to other economic grass crop species, where the comparative genome relationships are known in some detail. In this way, loci related to preharvest sprouting in wheat have been compared with some major genes affecting dormancy in maize and dormancy related QTLs in rice. This alignment identifies some candidate loci from maize and some regions of the rice genome that may relate to important wheat QTLs. In turn this approach will open up application of the emerging rice genomic DNA sequence to wheat pre-harvest sprouting research. This revised version was published online in August 2006 with corrections to the Cover Date.     

Spike-based and seed-based selection for preharvest sprouting resistance in wheat

Summary The utility of spike- and seed-based mass selection techniques for improving preharvest sprouting resistance in heterogeneous wheat (Triticum spp.) populations was evaluated. Sorting seed by size improved selection efficiency in some cases, putatively by physiological synchronization. Progeny testing, as well as changes in frequency of red-kernelled types, indicate effectiveness of both spike- and seed-based mass selection for reduced preharvest sprouting. Differential effectiveness of mass selection, in populations segregating for dormancy from different sources, is consistent with previous work on mechanisms of dormancy from these sources. These results are of value to improvement of preharvest sprouting resistance in large, heterogeneous wheat populations.     

Genetic and environmental control of dormancy in white-grained wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Grain dormancy in wheat is an important component of resistance to preharvest sprouting and hence an important trait for wheat breeders. The significant influence of environment on the dormancy phenotype makes this trait an obvious target for marker-assisted-selection. Closely related breeding lines, SUN325B and QT7475, containing a major dormancy QTL derived from AUS1408 located on chromosome 4A, but substantially different in dormancy phenotype, were compared with a non-dormant cultivar, Hartog, in a range of controlled environments. As temperature increased, dormancy at harvest-ripeness decreased particularly for QT7475. The dormancy phenotypes of reciprocal F₁ grains involving all possible combinations of Hartog, QT7475 and SUN325B were also compared in two environments with different temperatures. The results were consistent with the presence of QTL in addition to 4A in SUN325B, compared with QT7475, at least one of which was associated with the seed coat. Genetic analysis of a doubled haploid population derived from SUN325B × QT7475 identified a highly significant QTL located on chromosome 3BL, close to the expected position of the mutant allele of the red seed coat colour gene in white-grained wheat, R-B1a. When the lines in the population were grouped according to the parental alleles at marker loci flanking the 3B QTL, the dormancy phenotype frequency distribution for the SUN325B group was shifted towards greater dormancy compared with the QT7475 group. However, significant variation for dormancy phenotype remained within each group. Lines representing the extremes of the range of phenotypes within each group maintained their relative ranking across seven environments consistent with the presence of another unidentified QTL contributing to dormancy in SUN325B.     

Resistance to stripe rust in Triticum turgidum,T. tauschii and their synthetic hexaploids

Resistance to stripe rust (caused by Puccinia striiformis Westend.) of 34 Triticum turgidum L. var.durum, 278 T. tauschii, and 267 synthetic hexaploid wheats (T. turgidum x T. tauschii) was evaluated at the seedling stage in the greenhouse and at the adult-plant stage at two field locations. Mexican pathotype 14E14 was used in all studies. Seedling resistance, expressed as low infection type, was present in all three species. One hundred and twenty-eight (46%) accessions of T. tauschii, 8 (23%) of T. turgidum and 31 (12%) of synthetic hexaploid wheats were highly resistant as seedlings. In the field tests, resistance was evaluated by estimating area under disease progress curve (AUDPC). Synthetic hexaploid wheats showed a wide range of variability for disease responses in both greenhouse and field tests, indicating the presence of a number of genes for resistance. In general, genotypes with seedling resistance were also found to be resistant as adult plants. Genotypes, which were susceptible or intermediate as seedlings but resistant as adult plants, were present in both T. turgidum and the synthetic hexaploids. Resistances from either T. turgidum or T. tauschii or both were identified in the synthetic hexaploids in this study. These new sources of resistance could be incorporated into cultivated hexaploid wheats to increase the existing gene pool of resistance to stripe rust.     

On the problem of pre-harvest sprouting of wheat

Summary The problem of pre-harvest sprouting in wheat is very intricate. New concepts in breeding for resistance to pre-harvest sprouting regard many components such as -amylase production potential, response to gibberellic acid and inhibition in the bracts as important as dormancy. In the research work carried out at The University of Sydney's Plant Breeding Institute, we found varietal differences for each of these components. We also developed criteria for screening breeding material for these components. The implications of our findings in planning efficient breeding programmes to evolve varieties with multiple resistance to pre-harvest sprouting have been discussed briefly.     

Mapping genes for resistance to sprouting damage in wheat

A series of experiments to investigate the genetic basis of pre-harvest sprouting are reported. The results are combined with previously published studies in a composite genetic map for sprout resistance in hexaploid wheat. Different studies, using classical genetics, aneuploids, chromosome substitutions, or QTL mapping, have identified various regions of the A, B, and D genomes affecting dormancy. Comparisons between the available studies lead to the following conclusions: • Different studies often identify different genetic loci, in part reflecting different sampling from the available gene pool. This implies that many loci are involved in determining resistance, and that new loci may be discovered as the number of mapping studies increases.• There are, however, examples where similar map locations are implicated over different crosses. These may reflect the detection of key resistance genes. • Finally, (genotype × environment) interactions are frequently observed. The implications of these observations for crop improvement and research programmes are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Mapping and validation of PCR-based markers associated with a major QTL for seed dormancy in wheat

Seed dormancy is one of the important factors controlling pre-harvest sprouting (PHS) resistance in wheat. We identified a major quantitative trait locus (QTL) for seed dormancy on the long arm of wheat chromosome 4A (4AL) via simple sequence repeat (SSR)-based genetic mapping using doubled haploid lines from a cross between Japanese PHS resistant variety ‘Kitamoe’ and the Alpine non-resistant variety “Münstertaler” (K/M). The QTL explained 43.3% of total phenotypic variation for seed dormancy under greenhouse conditions. SSR markers flanking the QTL were assigned to the chromosome long arm fraction length 0.59–0.66 on the basis of chromosome deletion analysis, suggesting that the gene(s) controlling seed dormancy are probably located within this region. Under greenhouse conditions, the QTL explained 28.5 and 39.0% of total phenotypic variation for seed dormancy in Haruyutaka/Leader (HT/L) and OS21-5/Haruyokoi (O/HK) populations, respectively. However, in field conditions, the effect was relatively low or not significant in both the K/M and HT/L populations. These markers were considered to be widely useful in common with various genetic backgrounds for improvement of seed dormancy through the use of marker-assisted selection. Further detailed research using near isogenic lines will be needed to define how this major QTL interacts with environmental conditions in our area.     

Suppression/expression of resistance to stripe rust in synthetic hexaploid wheat (Triticum turgidum×T. tauschii)

Seventy-four hexaploid wheats, synthesized by either crossing resistantTriticum turgidum L. var.durum with susceptible/intermediateT. tauschii or susceptible/intermediateT. turgidum with resistantT. tauschii, and their parents were evaluated as seedlings in the greenhouse and as adult-plants at two field locations in Mexico for resistance to pathotype 14E14 of stripe (or yellow) rust (caused byPuccinia striiformis Westend). The seedlings of different synthetic hexaploids showed high phenotypic diversity for resistance. However, the resistance level of only 15 of the 74 synthetic hexaploid wheats were similar to the low infection types of the respective donor parents. The remaining synthetic wheats displayed either intermediate or high infection types. A similar result was also obtained in field tests, where only 18 synthetic hexaploids were resistant as adult-plants. In general, genotypes with seedling resistance were also resistant as adult-plants. A few synthetic hexaploids, which displayed intermediate or susceptible infection types as seedlings were resistant as adult-plants, indicating that additional genes for adult-plant resistance were also present. The fact that resistance of some donor parents was not expressed, or only partially expressed, in a synthetic hexaploid background suggests the presence of suppressor genes in the both the A or B, and D genomes ofT. turgidum andT. tauschii, respectively. The resistance of a donor parent was expressed in a synthetic hexaploid only if the corresponding suppressor was absent in the second parent. Moreover, the suppressors appeared to be resistance gene specific.     

Genetic analysis of resistance to root-lesion nematode (Pratylenchus thornei) in wheat

The inheritance of resistance to root‐lesion nematode was investigated in five synthetic hexaploid wheat lines and two bread wheat lines using a half‐diallel design of F₁ and F₂ crosses. The combining ability of resistance genes in the synthetic hexaploid wheat lines was compared with the performance of the bread wheat line ‘GS50a’, the source of resistance to Pratylenchus thornei used in Australian wheat breeding programmes. Replicated glasshouse trials identified P. thornei resistance as polygenic and additive in gene action. General combining ability (GCA) of the parents was more important than specific combining ability (SCA) effects in the inheritance of P. thornei resistance in both F₁ and F₂ populations. The synthetic hexaploid wheat line ‘CPI133872’ was identified as the best general combiner, however, all five synthetic hexaploid wheat lines possessed better GCA than ‘GS50a’ The synthetic hexaploid wheat lines contain novel sources of P. thornei resistance that will provide alternative and more effective sources of resistance to be utilized in wheat breeding programmes.     

Seed development-related expression of ARGONAUTE4_9 class of genes in barley: possible role in seed dormancy

Resistance to pre-harvest sprouting is an important breeding objective for cereal crops like barley and wheat. Seed dormancy, which determines the resistance or susceptibility to pre-harvest sprouting (PHS), is a complex trait. It is largely controlled by the antagonistic action of the plant hormones abscisic acid and gibberellic acid, but also has a large component of genotype?×?environment interaction. Recent studies have revealed a role for epigenetic changes through histone modification in controlling seed dormancy. However, the role of DNA methylation in seed development and dormancy is not known. In this study, we explored the role of ARGONAUTE4_9 class genes of the DNA methylation pathway in seed development and dormancy in barley. Our results show that the two AGO4_9 class genes in barley, i.e. AGO1002 and AGO1003, are preferentially expressed in ovaries at meiosis and in embryos 25?days after pollination (DAP). The expression of AGO1003 is two to fivefold higher than that of AGO1002 in these tissues, demonstrating differential expression of these genes. We also analysed the expression of AGO1003 in embryos of PHS-resistant and -susceptible varieties at 25?DAP and found a significant variation in the expression of this gene in seeds of dormant and non-dormant lines. The observed expression pattern of AGO1002 and AGO1003 suggests a possible role in sporogenesis and post-fertilization seed development. Indirectly these results imply a potential role of DNA methylation in seed development and seed dormancy.