长江流域小麦地方品种农艺品质性状的遗传多样性
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摘要
中国长江流域地域辽阔,地势复杂,气候多样,各生态区形成有种类丰富的小麦地方品种,存在着巨大的遗传变异。但截至目前,对整个长江流域小麦地方品种农艺、品质性状的遗传变异还缺乏较为系统的分析,只有对某些局部地区或部分小麦品种农艺、品质性状遗传多样性的研究报道,而对湖北小麦地方品种主要性状的报道较少,对长江流域小麦地方品种HMW-GS的MALDI-TOF-MS和对湖北小麦地方品种HMW-GS的SDS-PAGE分析未见报道。因此,为发掘适合湖北小麦的有利性状和可供小麦育种利用的优异遗传资源,也为小麦遗传多样性研究、种质资源研究、小麦遗传育种研究和优质小麦生产以及优良种质的开发利用打下基础和提供依据,以长江流域的485份小麦地方品种为材料,通过田间试验和实验室分析,对长江流域的小麦地方品种种质资源进行了农艺、品质性状多样性分析,在此基础上,对供试材料进行了HMW-GS的SDS-PAGE分析和MALDI-TOF-MS分析,结果如下：
1.研究中长江流域小麦地方品种供试材料农艺性状表现参差不齐,总体表现为高秆,早熟型,短穗型,小穗数、穗粒数偏少,千粒重较高,多数材料茎秆较细弱,穗部性状较差,籽粒偏小,易倒伏,为直接利用带来一定困难。各性状中穗粒重的变异程度最大,最不稳定,可选择范围大,其次是穗粒数和主穗穗颈至旗叶枕距,株高、抽穗期和小穗数的差异相对较小,表明其稳定性较好。虽然长江流域小麦地方品种农艺性状表现变异丰富,没有综合性状特别优异的材料,但仍然可以筛选到一些单一性状优良的材料。通过品种间的杂交,这些优良材料可直接应用于小麦的遗传改良。
在湖北种植的8个地区地方品种的农艺性状表现表明,除了主穗长、主穗小穗数、穗粒数外在其他性状上各个生态区间存在显著或极显著差异。每个生态区内都普遍存在丰富的变异,湖北省地方品种株高平均值均高于青藏高原和澳大利亚材料,都在120cm以上；青藏高原材料穗粒数、穗粒重、千粒重较大,澳大利亚材料穗粒数、穗粒重变异系数最小,穗粒重、千粒重最大。这些结果表明青藏高原和澳大利亚材料在湖北种植的农艺性状表现较好,可作为湖北小麦选育材料；湖北省各个生态区材料都有一定缺陷,需要合理利用,即可为育种提供优异资源。
通过简单相关分析,表明长江流域小麦地方品种各性状间存在着许多有利和不利的相关性。穗粒重、穗粒数和千粒重之间的显著相关说明穗粒数、千粒重的增加均有利于穗粒重的提高,但穗粒数和千粒重之间存在矛盾,穗粒数的增加会导致千粒重的降低,这与波兰小麦、西藏小麦、东方小麦、马卡小麦的研究结果相似。同时株高、主穗长与千粒重的负相关表明在长江流域小麦地方品种中株高和穗长的增加并不能引起千粒重的增加,反而使千粒重降低,这与王亚娟等261份来自全国各地优质小麦的农艺性状的相关分析结果相似,但与波兰小麦、波斯小麦、密穗小麦、东方小麦的研究结果不一致。长江流域小麦地方品种各性状间复杂的相关性表明在选择利用这些地方品种时中关注目标性状的同时也要兼顾其他性状。
2.长江流域小麦地方品种有丰富的中筋小麦种质资源,兼有强筋、弱筋小麦地方种质资源,与《专用小麦优势区域发展规划(2003-2007)》中的要求一致。长江流域小麦地方品种蛋白质含量达到强筋小麦品质指标(≥14.00%)占14.14%,达到中筋小麦品质指标(14.00%-11.50%)占77.85%,达到弱筋小麦品质指标(≤11.50%)占8.01%。长江流域小麦地方品种湿面筋达到强筋小麦品质指标(≥32%)占0.21%,达到弱筋小麦品质指标(32.00%-22.00%)占99.58%,达到弱筋小麦品质指标(≤22.00%)占0.21%。长江流域小麦地方品种沉降值达到强筋小麦指标(≥45.00mL)占10.97%,达到中筋小麦品质指标(45.00-30.OmL)占88.82%,达到弱筋小麦品质指标(≤30.OmL)占0.21%。因此,长江流域小麦地方品种是该区域小麦品质育种不可多得的种质资源,有极大的利用和开发价值。
对长江流域小麦地方品种品质性状相关性分析可知,淀粉含量与蛋白质含量、湿面筋、沉降值之间呈极显著负相关,表明淀粉含量对小麦品质有极大的影响,所以淀粉含量可以作为小麦品质的一个重要的品质指标,淀粉品质与蛋白质品质、湿面筋、沉降值之间存在密切联系,其共同影响着小麦的加工品质,在全面评价小麦品质或判断某一性状改变对加工品质的影响时,或根据不同的育种目标改良小麦的品质时,应综合考虑。湿面筋和蛋白质含量的相关系数为0.93**,相关系数高,说明湿面筋和蛋白质指标间协调一致,可作为小麦品质考察指标。
青藏高原地区小麦地方品种蛋白质含量、湿面筋含量、沉降值最高,与其他生态区差异达到极显著,鄂东南、鄂东、江汉平原、鄂西、鄂北、鄂西北生态区小麦地方品种之间蛋白质含量差异不显著。青藏高原地区小麦地方品种淀粉含量最低,与其他生态区差异达到极显著,鄂东南、鄂东、江汉平原、鄂西、鄂北、鄂西北生态区小麦地方品种之间淀粉含量差异不显著。表明青藏高原地区小麦地方品种在蛋白质含量、湿面筋含量、沉降值上优于湖北地区地方品种,且在湖北作冬小麦种植绝大部分品种均能正常成熟结实,因此可作为改良品质的种质资源加以利用。
3.供试材料中,亚基的分布范围较广,大部分材料含有7+8和2+12亚基,含5+10亚基的材料也较多,有27个,占32.14%。这些材料高分子量麦谷蛋白亚基组合有多种类型,主要亚基组合可分为三个大类,第一类是含7+8和2+12亚基的,第二类是含7+8和5+10亚基的,第三类是含未知亚基的。其中含第一类亚基的小麦材料有286个,占60.47%；含第二类亚基的材料有26个,占5.50%；含第三类亚基的材料有50个,占10.57%。
供试材料在编码高分子量麦谷蛋白亚基的Glu-A1、Glu-B1和Glu-D1这3个基因位点上表现出丰富的多态性,共有22种HMW-GS等位变异。其中,Glu-A1位点上有3种等位变异类型；Glu-B1位点检测到的等位变异类型最多,为16种；Glu-D1位点有3种。从各个亚基的出现频率看,在Glu-A1位点上的3种变异类型0、1、2*中,0亚基频率最高,为89.22%,1亚基次之,为9.73%,2*亚基最少,为1.05%。Glu-B1位点上的亚基类型最为丰富,检测到的16种亚基类型中,7+8亚基出现频率最高,为65.96%；其次为?+8亚基,为9.51%；13+16、17+18、7、20亚基为稀有亚基类型,频率在1.69%～3.59%。在Glu-D1位点上,2+12亚基类型最多,频率为89.64%,其次是优质亚基5+10,占9.30%。
通过对供试材料的HMW-GS组合形式和频率研究,可以发现供试材料的HMW-GS有44种组合形式。每种形式的亚基组合一般包含3至6个亚基。供试的473份材料各亚基组合形式较为分散,其中(0,7+8,2+12)组合频率最高,为54.97%。其次为(0,?+8,2+12)、(1,7±8,2+12)、(0,7+8,5+10)、(0,7+9,2+12)4种组合形式,频率依次为8.46%、5.07%、4.23%、4.23%。其它为稀有亚基组合形式,频率在0.21%～2.96%。供试材料中,Glu-A1位点的优质亚基为51个,占18.68%；Glu-B1位点优质亚基为73个,占15.43%；Glu-D1位点优质亚基为44个,占16.12%。Glu-1的3个位点均具有优质亚基的材料8个,占1.69%；仅2个位点均具有优质亚基的材料51个,占10.78%；只有1个位点具有优质亚基的材料351个,占74.21%。
在供试的材料中,虽然优质亚基组合所占比例不高,但优质亚基占有较高的比例,尤其是出现了50个未知亚基类型。这些材料将是小麦品质遗传育种的宝贵资源。对这些未知亚基开展进一步研究,对他们予以命名和弄清其对小麦品质的贡献,将对小麦品质育种具有极其重要的意义。
4.MALDI-TOF-MS分析发现,大量的高分子量谷蛋白亚基等位基因变异在所研究的地方品种中。其中453份材料的高分子量麦谷蛋白亚基组成是同质的,32份异质的,37份含有异常亚基。共有22个正常的Glu-1位点等位基因被检测出来,3个属于Glu-A1,13个属于Glu-B1,6个属于GIu-D1,分布于63种不同的等位基因组合中。
在研究中,除了正常的22个等位基因外,12个等位基因编码了分子量69100,69900,73000,73100,75140,75600,76800,79000,79100,79800,83200,84300的异常亚基也被检测到。这代表了异常的等位基因比以前的报告有一个更高的速率。研究还发现,大量的品种中有一个或两个HMWGS基因沉默,特别是16个品种中1Bx的基因沉默。这些沉默的等位基因是解析小麦品质中特定等位基因效应的宝贵资源。
总体来说,这项研究研究了485份中国的长江中上游地区小麦地方品种的高分子量谷蛋白亚基组成。在这项研究中获得的信息可用于小麦育种,培育新的品种来满足特定的最终产品的需求。在一般情况下,22个高分子量麦谷蛋白亚基等位基因和16个异常亚基在485份中国长江中上游地区收集到的地方品种中被鉴定出来。这些新的等位基因的进一步研究,目前正在获得其编码序列,将它们与先前报道的新的等位基因相比较。
Yangtze River basin is complex terrain with diversitifed climates. Abundant local wheat varieties have been developed in this area. The research on agronomic and quality characteristics has always been the key of crop improvement in agricultural production. Even when the molecular markers are developing fast these days, fully understanding and scientific assessment of the representing reality of the agronomic and quality characteristic of germplasm materials is still of great importance for rational use of germplasm resources and parent selection. Unfortunately, systematic analysis on the genetic variation of agronomic and quality characteristics of local wheat varieties in the Yangtze River basin is still lacking. The previous researches mainly focused on the genetic diversity of agronomic and quality characteristics of regional or partial varieties. No study was conducted on the analysis of High-molecular-weight glutenin subunits(HMW-GS) of local varieties in Hubei province. We characterized genetic diversity and HMW-GS in477wheat varieties from the Yangzi River basin. using agronomic and quality characteristics. The major findings of SDS-PAGE analysis and MALDI-TOF-MS analysis on the HMW-GS are reported as follows.
1. Generally speaking, the wheat varieties in Yangzi River basin studied showed high stalk, precocity, short head, small spikelet number and grain number per spike, high kernel weight. The majority of them have weak stem, poor ear characters, small grain, and are susceptible to lodging. Among the characters studied, spike grain weigh showed wide range of variation followed by the grain number per spike, and the distance from main spike neck to flag leaf pulvinus. No significantly differences were found for plant height, heading date and the number of spikelet number among the varieties.
The agronomic traits studied showed significantly difference among different ecological zones and within each zone except for main panicle length, spike numbers of main spike, grain number per spike The averaged plant height of Hubei varieties, which is above120cm, is higher than that of materials from Tibetan Plateau and Australia. The grain number per spike, spike grain weight and kernel weigh in Tibetan varieties are than those in the varieties from Yangtz river region. For the varieties from Australia, The grain number per spike and spike grain weight showed the lowest variation, and the spike grain weight and kernel weigh showed the highest variation among the varieties studied. The results indicated that the agronomic characters of wheat materials from Tibetan Plateau and Australia perform well in Hubei, thus they could be used for wheat breeding. The varieties from the ecological zones in Hubei displyed some undisrable chatacters,it needs better strategy to eccifcetly use the germplasm from Hubei for breeding program.
Correlation analysis found significant correlation between grain number per spike, spike grain weight and kernel weigh, indicating that the increase of grain number per spike and kernel weight could redult in the increase of spike grain weight. However, the grain number per spike was nagetively correlated with kernel weigh, increase of grain number per spike could cause the decrease of kernel weight, which is similar to the reports on Polish wheat, Tibet wheat, orient wheat and macha wheat. The negative correlation between plant height, main panicle length and kernel weight indicated that the increase of plant height and panicle length would decrease the kernel weight, which is consistent with the correlation analysis on the261wheat materials collected from all over the country; but is inconsistent with the study on Polish wheat, Tibet wheat, orient wheat and macha wheat.
2. The results indicated that the wheat landraces from Yangtze River valley were rich in medium-gluten, few with strong gluten and weak gluten, which was consistent with proposal made in the 《Special-purpose Wheat Regional Development Plan (2003-2007)》.14.14%,77.85%and8.01%of wheat landraces are with protein content reaching high strength gluten index (≥14.00%), middle strength gluten index(14.00%-11.50%), low strength gluten index (≤11.50%) p, respectively. Wheat landraces of wet gluten content reaching high-strength-gluten index (≥32%), middle-strength-gluten index (32.00%～22.00%),low-strength-gluten index (≤22.00%) accouted for0.21%,99.58%,0.21%, respectively. The percentage of local varieties with zeleny reaching high-strength-gluten index (≥45.00mL), middle-strength-gluten index (45.00-30.0mL), low-strength-gluten index (≤30.0mL) were10.97%,88.82%,0.21%, respectively. Therefore, landraces in Yangtze River region of China are valuable resources in wheat breeding.
The correlation analysis indicated that starch content was significantly negatively correlated with protein content, wet gluten content, zeleny Starch quality was closely associated with protein quality, wet gluten content, zeleny. These four components determine wheat processing quality. The wet gluten content showed a highly positive correlation with the protein content, with correlation coefficient of0.93.. The wet gluten content can be used as a quality index to evaluate wheat quality.
The protein content, wet gluten content, zeleny in the Tibetan local varieties of wheat were significantly higher than those invarieties from other ecological areas. Both protein and starch content did not showed significantly difference among landraces from South-eastern Hubei, East of Hubei, Jianghan plain, West of Hubei, North of Hubei, North-west of Hubei.The starch content of Tibetan local varieties was significantly lower than that in samples from other ecological areas. The results demonstrated that protein content, wet gluten content, zeleny of landraces from Qingzang plateau were superior to those in the local varieties from Hubei Province. Most of the landraces from Qingzang plateau planted in winter in Hubei could normally mature and yield. Therefore they are useful germplasm to improve the wheat quality.
3. Among the varieties tested,7+8and2+12subunits are detected in most varieties; and5+10subunit is also very common, and32.14%varieties possessed this subunit. The combinations of high molecular weight glutenin subunits in these varieties tested were mainly divided into3types.286(60.4%) varities contained7+8and2+12subunits,26(5.50%) varieties contained7+8and5+10subunits, and50(10.57%) varieties contained undefined subunits.A total of22alleles are found among the varieties tested, with3alleles for Glu-Al;16alleles for Glu-Bl; and3alleles for Glu-D1locus. Of the3alleles on Glu-A1,0,1,2*, the frequency of0subunit is the highest (89.22%), followed by1subunit (9.73%); and the frequency of2*subunit(1.05%) is the lowest one. Of the16detected subunit types (alleles) at Glu B1,65.96%are7+8subunit followed by?+8subunit (9.51%); the13+16,17+18,7, and20subunits are rare types, with their frequency ranking from1.69%to3.59%. At Glu-Dl site,2+12subunit type is the most common (89.64%), followed by high quality subunit5+10(9.30%).
A total of44HMW-GS combination forms are revealed in the varieties. Generally, each combination is consist of3-6subunits. The frequency of subunit combination forms detected in473varieties are quite difference from each other, among which combination form (0,7+8,2+12) is the most frequent(54.97%), followed by (0,?+8,2+12)(8.46%),(1,7+8,2+12)(5.07%),(0,7+8,5+10)(4.23%) and (0,7+9,2+12)(4.23%); the rest are rare subunit combinations, with frequency of0.21%～2.96%. The high quality subunits at Glu-A1site were detected in51varieties (18.68%); at Glu-B1site were detected in73varieties (15.43%); and at Glu-D1site in44varieties (16.12%). Eight varieties have high quality subunits at all the3Glu-1sites, accounting for1.69%of the total;518materials have high quality subunits at2of the3Glu-1sites, accounting for10.78%of the total;351materials have high quality subunits at1of the3Glu-1sites, accounting for74.21%of the total.
Although the proportion of high quality subunit combinations is not high among the studied materials, the ratio of high quality subunits is relatively high; especially50unidentified subunit materials are found. They are precious resource for genetic breeding of wheat. Future study on these materials to reveal their contribution to the wheat quality, is of very importance to wheat quality breeding.
4. Extensive allelic variation in HMW glutenin subunits was detected among the studied landraces. A total of453of the accessions were homogeneous for HMW-GS composition,32were heterogeneous and37accessions contained abnormal subunits. A total of22normal alleles for the Glu-1loci were detected,3belonged to Glu-A1,13at Glu-B1and6at Glu-D1, resulting in63different allele combinations.
In our study, apart from the22normal alleles, twelve alleles encoding abnormal subunits with molecular weights of69,100,69,900,73,000,73,100,75,140,75,600,76,800,79,000,79,100,79,800,83,200,84,300were also detected. This represented a higher rate of abnormal alleles than previous reports, which is likely due to the enhanced resolution in determining the molecular mass of the MALDI-TOF technology. In this study, a large numer of lines had one or two HMWGS genes silenced, especially16lines with the1Bx gene silenced, of which the silencing mechanism is still unclear. These null alleles are valuable resources for dissecting specific allele effects in wheat quality.
To conclude, this study characterized the HMW glutenin subunit compositions of485wheat landraces in the Yangtze-River region of China. The information obtained in this study may be used by wheat breeders for breeding new cultivars to meet specific end-product needs. In general,22HMW-GS alleles with16abnormal subunits were identified in a collection of485landraces from the Yangtze-River region of China. Further studies of these novel alleles are currently undergoing to obtain their coding sequences in order to match them with previously reported novel alleles.
1.研究中长江流域小麦地方品种供试材料农艺性状表现参差不齐,总体表现为高秆,早熟型,短穗型,小穗数、穗粒数偏少,千粒重较高,多数材料茎秆较细弱,穗部性状较差,籽粒偏小,易倒伏,为直接利用带来一定困难。各性状中穗粒重的变异程度最大,最不稳定,可选择范围大,其次是穗粒数和主穗穗颈至旗叶枕距,株高、抽穗期和小穗数的差异相对较小,表明其稳定性较好。虽然长江流域小麦地方品种农艺性状表现变异丰富,没有综合性状特别优异的材料,但仍然可以筛选到一些单一性状优良的材料。通过品种间的杂交,这些优良材料可直接应用于小麦的遗传改良。
在湖北种植的8个地区地方品种的农艺性状表现表明,除了主穗长、主穗小穗数、穗粒数外在其他性状上各个生态区间存在显著或极显著差异。每个生态区内都普遍存在丰富的变异,湖北省地方品种株高平均值均高于青藏高原和澳大利亚材料,都在120cm以上；青藏高原材料穗粒数、穗粒重、千粒重较大,澳大利亚材料穗粒数、穗粒重变异系数最小,穗粒重、千粒重最大。这些结果表明青藏高原和澳大利亚材料在湖北种植的农艺性状表现较好,可作为湖北小麦选育材料；湖北省各个生态区材料都有一定缺陷,需要合理利用,即可为育种提供优异资源。
通过简单相关分析,表明长江流域小麦地方品种各性状间存在着许多有利和不利的相关性。穗粒重、穗粒数和千粒重之间的显著相关说明穗粒数、千粒重的增加均有利于穗粒重的提高,但穗粒数和千粒重之间存在矛盾,穗粒数的增加会导致千粒重的降低,这与波兰小麦、西藏小麦、东方小麦、马卡小麦的研究结果相似。同时株高、主穗长与千粒重的负相关表明在长江流域小麦地方品种中株高和穗长的增加并不能引起千粒重的增加,反而使千粒重降低,这与王亚娟等261份来自全国各地优质小麦的农艺性状的相关分析结果相似,但与波兰小麦、波斯小麦、密穗小麦、东方小麦的研究结果不一致。长江流域小麦地方品种各性状间复杂的相关性表明在选择利用这些地方品种时中关注目标性状的同时也要兼顾其他性状。
2.长江流域小麦地方品种有丰富的中筋小麦种质资源,兼有强筋、弱筋小麦地方种质资源,与《专用小麦优势区域发展规划(2003-2007)》中的要求一致。长江流域小麦地方品种蛋白质含量达到强筋小麦品质指标(≥14.00%)占14.14%,达到中筋小麦品质指标(14.00%-11.50%)占77.85%,达到弱筋小麦品质指标(≤11.50%)占8.01%。长江流域小麦地方品种湿面筋达到强筋小麦品质指标(≥32%)占0.21%,达到弱筋小麦品质指标(32.00%-22.00%)占99.58%,达到弱筋小麦品质指标(≤22.00%)占0.21%。长江流域小麦地方品种沉降值达到强筋小麦指标(≥45.00mL)占10.97%,达到中筋小麦品质指标(45.00-30.OmL)占88.82%,达到弱筋小麦品质指标(≤30.OmL)占0.21%。因此,长江流域小麦地方品种是该区域小麦品质育种不可多得的种质资源,有极大的利用和开发价值。
对长江流域小麦地方品种品质性状相关性分析可知,淀粉含量与蛋白质含量、湿面筋、沉降值之间呈极显著负相关,表明淀粉含量对小麦品质有极大的影响,所以淀粉含量可以作为小麦品质的一个重要的品质指标,淀粉品质与蛋白质品质、湿面筋、沉降值之间存在密切联系,其共同影响着小麦的加工品质,在全面评价小麦品质或判断某一性状改变对加工品质的影响时,或根据不同的育种目标改良小麦的品质时,应综合考虑。湿面筋和蛋白质含量的相关系数为0.93**,相关系数高,说明湿面筋和蛋白质指标间协调一致,可作为小麦品质考察指标。
青藏高原地区小麦地方品种蛋白质含量、湿面筋含量、沉降值最高,与其他生态区差异达到极显著,鄂东南、鄂东、江汉平原、鄂西、鄂北、鄂西北生态区小麦地方品种之间蛋白质含量差异不显著。青藏高原地区小麦地方品种淀粉含量最低,与其他生态区差异达到极显著,鄂东南、鄂东、江汉平原、鄂西、鄂北、鄂西北生态区小麦地方品种之间淀粉含量差异不显著。表明青藏高原地区小麦地方品种在蛋白质含量、湿面筋含量、沉降值上优于湖北地区地方品种,且在湖北作冬小麦种植绝大部分品种均能正常成熟结实,因此可作为改良品质的种质资源加以利用。
3.供试材料中,亚基的分布范围较广,大部分材料含有7+8和2+12亚基,含5+10亚基的材料也较多,有27个,占32.14%。这些材料高分子量麦谷蛋白亚基组合有多种类型,主要亚基组合可分为三个大类,第一类是含7+8和2+12亚基的,第二类是含7+8和5+10亚基的,第三类是含未知亚基的。其中含第一类亚基的小麦材料有286个,占60.47%；含第二类亚基的材料有26个,占5.50%；含第三类亚基的材料有50个,占10.57%。
供试材料在编码高分子量麦谷蛋白亚基的Glu-A1、Glu-B1和Glu-D1这3个基因位点上表现出丰富的多态性,共有22种HMW-GS等位变异。其中,Glu-A1位点上有3种等位变异类型；Glu-B1位点检测到的等位变异类型最多,为16种；Glu-D1位点有3种。从各个亚基的出现频率看,在Glu-A1位点上的3种变异类型0、1、2*中,0亚基频率最高,为89.22%,1亚基次之,为9.73%,2*亚基最少,为1.05%。Glu-B1位点上的亚基类型最为丰富,检测到的16种亚基类型中,7+8亚基出现频率最高,为65.96%；其次为?+8亚基,为9.51%；13+16、17+18、7、20亚基为稀有亚基类型,频率在1.69%～3.59%。在Glu-D1位点上,2+12亚基类型最多,频率为89.64%,其次是优质亚基5+10,占9.30%。
通过对供试材料的HMW-GS组合形式和频率研究,可以发现供试材料的HMW-GS有44种组合形式。每种形式的亚基组合一般包含3至6个亚基。供试的473份材料各亚基组合形式较为分散,其中(0,7+8,2+12)组合频率最高,为54.97%。其次为(0,?+8,2+12)、(1,7±8,2+12)、(0,7+8,5+10)、(0,7+9,2+12)4种组合形式,频率依次为8.46%、5.07%、4.23%、4.23%。其它为稀有亚基组合形式,频率在0.21%～2.96%。供试材料中,Glu-A1位点的优质亚基为51个,占18.68%；Glu-B1位点优质亚基为73个,占15.43%；Glu-D1位点优质亚基为44个,占16.12%。Glu-1的3个位点均具有优质亚基的材料8个,占1.69%；仅2个位点均具有优质亚基的材料51个,占10.78%；只有1个位点具有优质亚基的材料351个,占74.21%。
在供试的材料中,虽然优质亚基组合所占比例不高,但优质亚基占有较高的比例,尤其是出现了50个未知亚基类型。这些材料将是小麦品质遗传育种的宝贵资源。对这些未知亚基开展进一步研究,对他们予以命名和弄清其对小麦品质的贡献,将对小麦品质育种具有极其重要的意义。
4.MALDI-TOF-MS分析发现,大量的高分子量谷蛋白亚基等位基因变异在所研究的地方品种中。其中453份材料的高分子量麦谷蛋白亚基组成是同质的,32份异质的,37份含有异常亚基。共有22个正常的Glu-1位点等位基因被检测出来,3个属于Glu-A1,13个属于Glu-B1,6个属于GIu-D1,分布于63种不同的等位基因组合中。
在研究中,除了正常的22个等位基因外,12个等位基因编码了分子量69100,69900,73000,73100,75140,75600,76800,79000,79100,79800,83200,84300的异常亚基也被检测到。这代表了异常的等位基因比以前的报告有一个更高的速率。研究还发现,大量的品种中有一个或两个HMWGS基因沉默,特别是16个品种中1Bx的基因沉默。这些沉默的等位基因是解析小麦品质中特定等位基因效应的宝贵资源。
总体来说,这项研究研究了485份中国的长江中上游地区小麦地方品种的高分子量谷蛋白亚基组成。在这项研究中获得的信息可用于小麦育种,培育新的品种来满足特定的最终产品的需求。在一般情况下,22个高分子量麦谷蛋白亚基等位基因和16个异常亚基在485份中国长江中上游地区收集到的地方品种中被鉴定出来。这些新的等位基因的进一步研究,目前正在获得其编码序列,将它们与先前报道的新的等位基因相比较。
Yangtze River basin is complex terrain with diversitifed climates. Abundant local wheat varieties have been developed in this area. The research on agronomic and quality characteristics has always been the key of crop improvement in agricultural production. Even when the molecular markers are developing fast these days, fully understanding and scientific assessment of the representing reality of the agronomic and quality characteristic of germplasm materials is still of great importance for rational use of germplasm resources and parent selection. Unfortunately, systematic analysis on the genetic variation of agronomic and quality characteristics of local wheat varieties in the Yangtze River basin is still lacking. The previous researches mainly focused on the genetic diversity of agronomic and quality characteristics of regional or partial varieties. No study was conducted on the analysis of High-molecular-weight glutenin subunits(HMW-GS) of local varieties in Hubei province. We characterized genetic diversity and HMW-GS in477wheat varieties from the Yangzi River basin. using agronomic and quality characteristics. The major findings of SDS-PAGE analysis and MALDI-TOF-MS analysis on the HMW-GS are reported as follows.
1. Generally speaking, the wheat varieties in Yangzi River basin studied showed high stalk, precocity, short head, small spikelet number and grain number per spike, high kernel weight. The majority of them have weak stem, poor ear characters, small grain, and are susceptible to lodging. Among the characters studied, spike grain weigh showed wide range of variation followed by the grain number per spike, and the distance from main spike neck to flag leaf pulvinus. No significantly differences were found for plant height, heading date and the number of spikelet number among the varieties.
The agronomic traits studied showed significantly difference among different ecological zones and within each zone except for main panicle length, spike numbers of main spike, grain number per spike The averaged plant height of Hubei varieties, which is above120cm, is higher than that of materials from Tibetan Plateau and Australia. The grain number per spike, spike grain weight and kernel weigh in Tibetan varieties are than those in the varieties from Yangtz river region. For the varieties from Australia, The grain number per spike and spike grain weight showed the lowest variation, and the spike grain weight and kernel weigh showed the highest variation among the varieties studied. The results indicated that the agronomic characters of wheat materials from Tibetan Plateau and Australia perform well in Hubei, thus they could be used for wheat breeding. The varieties from the ecological zones in Hubei displyed some undisrable chatacters,it needs better strategy to eccifcetly use the germplasm from Hubei for breeding program.
Correlation analysis found significant correlation between grain number per spike, spike grain weight and kernel weigh, indicating that the increase of grain number per spike and kernel weight could redult in the increase of spike grain weight. However, the grain number per spike was nagetively correlated with kernel weigh, increase of grain number per spike could cause the decrease of kernel weight, which is similar to the reports on Polish wheat, Tibet wheat, orient wheat and macha wheat. The negative correlation between plant height, main panicle length and kernel weight indicated that the increase of plant height and panicle length would decrease the kernel weight, which is consistent with the correlation analysis on the261wheat materials collected from all over the country; but is inconsistent with the study on Polish wheat, Tibet wheat, orient wheat and macha wheat.
2. The results indicated that the wheat landraces from Yangtze River valley were rich in medium-gluten, few with strong gluten and weak gluten, which was consistent with proposal made in the 《Special-purpose Wheat Regional Development Plan (2003-2007)》.14.14%,77.85%and8.01%of wheat landraces are with protein content reaching high strength gluten index (≥14.00%), middle strength gluten index(14.00%-11.50%), low strength gluten index (≤11.50%) p, respectively. Wheat landraces of wet gluten content reaching high-strength-gluten index (≥32%), middle-strength-gluten index (32.00%～22.00%),low-strength-gluten index (≤22.00%) accouted for0.21%,99.58%,0.21%, respectively. The percentage of local varieties with zeleny reaching high-strength-gluten index (≥45.00mL), middle-strength-gluten index (45.00-30.0mL), low-strength-gluten index (≤30.0mL) were10.97%,88.82%,0.21%, respectively. Therefore, landraces in Yangtze River region of China are valuable resources in wheat breeding.
The correlation analysis indicated that starch content was significantly negatively correlated with protein content, wet gluten content, zeleny Starch quality was closely associated with protein quality, wet gluten content, zeleny. These four components determine wheat processing quality. The wet gluten content showed a highly positive correlation with the protein content, with correlation coefficient of0.93.. The wet gluten content can be used as a quality index to evaluate wheat quality.
The protein content, wet gluten content, zeleny in the Tibetan local varieties of wheat were significantly higher than those invarieties from other ecological areas. Both protein and starch content did not showed significantly difference among landraces from South-eastern Hubei, East of Hubei, Jianghan plain, West of Hubei, North of Hubei, North-west of Hubei.The starch content of Tibetan local varieties was significantly lower than that in samples from other ecological areas. The results demonstrated that protein content, wet gluten content, zeleny of landraces from Qingzang plateau were superior to those in the local varieties from Hubei Province. Most of the landraces from Qingzang plateau planted in winter in Hubei could normally mature and yield. Therefore they are useful germplasm to improve the wheat quality.
3. Among the varieties tested,7+8and2+12subunits are detected in most varieties; and5+10subunit is also very common, and32.14%varieties possessed this subunit. The combinations of high molecular weight glutenin subunits in these varieties tested were mainly divided into3types.286(60.4%) varities contained7+8and2+12subunits,26(5.50%) varieties contained7+8and5+10subunits, and50(10.57%) varieties contained undefined subunits.A total of22alleles are found among the varieties tested, with3alleles for Glu-Al;16alleles for Glu-Bl; and3alleles for Glu-D1locus. Of the3alleles on Glu-A1,0,1,2*, the frequency of0subunit is the highest (89.22%), followed by1subunit (9.73%); and the frequency of2*subunit(1.05%) is the lowest one. Of the16detected subunit types (alleles) at Glu B1,65.96%are7+8subunit followed by?+8subunit (9.51%); the13+16,17+18,7, and20subunits are rare types, with their frequency ranking from1.69%to3.59%. At Glu-Dl site,2+12subunit type is the most common (89.64%), followed by high quality subunit5+10(9.30%).
A total of44HMW-GS combination forms are revealed in the varieties. Generally, each combination is consist of3-6subunits. The frequency of subunit combination forms detected in473varieties are quite difference from each other, among which combination form (0,7+8,2+12) is the most frequent(54.97%), followed by (0,?+8,2+12)(8.46%),(1,7+8,2+12)(5.07%),(0,7+8,5+10)(4.23%) and (0,7+9,2+12)(4.23%); the rest are rare subunit combinations, with frequency of0.21%～2.96%. The high quality subunits at Glu-A1site were detected in51varieties (18.68%); at Glu-B1site were detected in73varieties (15.43%); and at Glu-D1site in44varieties (16.12%). Eight varieties have high quality subunits at all the3Glu-1sites, accounting for1.69%of the total;518materials have high quality subunits at2of the3Glu-1sites, accounting for10.78%of the total;351materials have high quality subunits at1of the3Glu-1sites, accounting for74.21%of the total.
Although the proportion of high quality subunit combinations is not high among the studied materials, the ratio of high quality subunits is relatively high; especially50unidentified subunit materials are found. They are precious resource for genetic breeding of wheat. Future study on these materials to reveal their contribution to the wheat quality, is of very importance to wheat quality breeding.
4. Extensive allelic variation in HMW glutenin subunits was detected among the studied landraces. A total of453of the accessions were homogeneous for HMW-GS composition,32were heterogeneous and37accessions contained abnormal subunits. A total of22normal alleles for the Glu-1loci were detected,3belonged to Glu-A1,13at Glu-B1and6at Glu-D1, resulting in63different allele combinations.
In our study, apart from the22normal alleles, twelve alleles encoding abnormal subunits with molecular weights of69,100,69,900,73,000,73,100,75,140,75,600,76,800,79,000,79,100,79,800,83,200,84,300were also detected. This represented a higher rate of abnormal alleles than previous reports, which is likely due to the enhanced resolution in determining the molecular mass of the MALDI-TOF technology. In this study, a large numer of lines had one or two HMWGS genes silenced, especially16lines with the1Bx gene silenced, of which the silencing mechanism is still unclear. These null alleles are valuable resources for dissecting specific allele effects in wheat quality.
To conclude, this study characterized the HMW glutenin subunit compositions of485wheat landraces in the Yangtze-River region of China. The information obtained in this study may be used by wheat breeders for breeding new cultivars to meet specific end-product needs. In general,22HMW-GS alleles with16abnormal subunits were identified in a collection of485landraces from the Yangtze-River region of China. Further studies of these novel alleles are currently undergoing to obtain their coding sequences in order to match them with previously reported novel alleles.
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