摘要
东北地区和黄淮海流域是我国两大大豆主产区，总产量占全国的3/4以上，迄今育成大豆品种的数量有1000多个，而得到大面积种植的只有100多个，约占10%。本研究于2009、2010和2011年在东北和黄淮海地区选择12个地点种植该区域自20世纪40年代以来大面积种植的89个大豆品种，调查产量构成因子、植株形态性状和生育期等重要性状，测定籽粒品质性状，主要包括粗蛋白、粗脂肪、5种脂肪酸、碳水化合物、维生素E、磷脂、微量元素、类胡萝卜素及异黄酮、皂苷、凝集素和包曼-伯克胰蛋白酶抑制因子等品质成分，对其中的63个品种进行了光、温敏感度鉴定。此外，利用品种的系谱关系，对113份大面积种植品种的祖先亲本进行了分析；选择与产量、株高、生育期及品质等性状相关的SSR标记，对89个品种进行了遗传多样性分析，并结合表型数据进行了关联分析。主要试验结果如下：
1.农艺性状的演变趋势
东北和黄淮海地区品种性状演变的整体趋势是：随育成年代推进，大豆品种的生育前期缩短、后期延长，株高降低，有效分枝数减少，抗倒伏能力显著增强，主茎节间缩短，产量显著升高。其中，黑龙江北部品种产量的提高主要依靠单株荚数和粒数增加；黑龙江中南部品种主要依靠荚粒数增多，荚数和百粒重变化不大；吉林-辽宁地区品种主要依靠百粒重的提高，单株荚数和粒数变化很小；黄淮海地区品种依靠百粒重的提高，而单株荚数和粒数随年代推进显著减少。在4个地区中，吉林-辽宁地区大面积种植品种的性状变化最小。
2.品质性状的演变趋势
供试品种粗蛋白和总异黄酮含量从北到南递增，东北地区品种的粗脂肪、总维生素E、总类胡萝卜素含量明显高于黄淮海地区品种。黑龙江北部品种亚油酸、亚麻酸、粗淀粉、低聚糖、总异黄酮和凝集素的含量随品种育成年代推进显著增加；黑龙江中南部品种亚油酸、棕榈酸和总异黄酮的含量随品种育成年代推进显著增加；吉林-辽宁地区品种粗淀粉含量随品种育成年代推进显著升高；黄淮海地区品种的油酸、硬脂酸和凝集素的含量随品种育成年代推进显著增加。其他品质性状变化较小。
3.光温反应性状的演变趋势
黑龙江北部和黄淮海地区品种在春播和夏播条件下的PS（光周期敏感度）随育成年度推进显著下降，黑龙江中南部品种在夏播条件下的PS显著降低；黑龙江中南部和黄淮海地区品种在短日条件下的TS（温度敏感度）显著下降，而在长日照条件下，黄淮海地区品种TS显著上升；黄淮海地区品种ΔPS（不同温度下的光周期敏感度）、ΔTS（不同光照下的温度敏感度）和PTCS（光温综合敏感度）显著下降。上述地区大面积种植品种的PS、TS及PTCS表现出随纬度降低逐渐增大的趋势，而ΔPS和ΔTS则随着纬度的降低逐渐减小。
4.大面积种植品种的系谱分析
通过系谱分析，归纳出大面积种植品种的直接亲本和祖先亲本。结果表明，大面积种植品种比普通品种拥有更多的祖先亲本；新近育成的品种比早期品种拥有更多的祖先亲本；黄淮海流域品种比东北地区品种拥有更多的祖先亲本。作者提出，以大面积种植品种作为“平台亲本”培育的新品种更容易在生产上种植应用，更有希望成为新的大面积种植品种。
5.大面积种植品种的遗传多样性分析
利用与大豆产量、品质、抗逆性、适应性等重要性状相关的125对SSR引物对89个大豆品种进行遗传多样性分析。结果表明，自北向南品种SSR标记的多态性呈逐渐升高的趋势，黑龙江北部、黑龙江中南部、吉林-辽宁地区和黄淮海地区品种SSR标记的多态性信息含量（PIC）依次为0.414、0.469、0.522和0.562。除黑龙江北部外，其余3个地区20世纪80年代以来大面积种植品种的多态性信息含量均比80年代以前种植的品种高。根据Nei和Li提出的遗传相似系数对供试品种进行聚类分析，发现除黄淮海北部地区的冀豆12外，东北和黄淮海地区品种各自聚成一类，而在东北地区品种中，黑龙江和吉林-辽宁地区的品种归属不同的亚类。
6.大面积种植品种优异等位变异发掘
在本研究检测的SSR位点中，共有141个位点（次）的变异与24个性状的变异关联，其中有39个位点（次）的变异与产量、荚数、粒数和百粒重等6个产量性状的变异关联，有19个位点的变异与株高、抗倒伏性等形态性状和成熟期等发育性状的变异关联，有82个位点（次）的变异与粗蛋白、脂肪酸、低聚糖、类胡萝卜素、胰蛋白酶抑制因子（BBI）、凝集素和总异黄酮含量等品质性状的变异关联。分析标记对性状的表型解释率、增效最大位点和减效最大位点及其典型载体材料，发现表型值大的品种含有更多的增效等位位点，表型值小的品种含有更多的减效等位位点。
The Northeast and Yellow-Huai-Hai Rivers Valley are the top two soybean growing regions inChina. The annual soybean production in the above two regions occupied above three quarters of thetotal output in China. Although over1000soybean varieties were released in the two regions in thepast century, only approximate10percent of these varieties were widely planted. To identify the yieldrelated charcters, plant morphological and growth period traits,89soybean varieties released since1940s were grown in12sites located in Northeast and Yellow-Huai-Hai Rivers Valley during theyears of2009,2010and2011, respectively. In addition,63varieties were selected to evaluate thephoto-thermal responses, and125SSR markers related with yield, plant height, growth period, andquality were selected to identify genetic diversity of89soybean varieties. Furthermore, to combinedata mentioned about with related phenotype data, association mapping were operated to search forthe elite alleles in these varieties. The main results are as follows:
1. Agronomic traits evolution of widely-planted varieties originated from different regions anddecades.
The overall trait evolution trends of varieties were summarized as: the days to flowering shortedand the days from flowering to maturity prolonged; the plant height and branche number per plantdeclined; lodging resistance enhanced and yield increased. The yield increasement of varieties fromthe northern Heilongjiang was due to increased numbers of pods and seeds per plant, while that ofvarieties from the middle and south parts of Heilongjiang attributed to increased seed numbers perpod, however, the pod number per plant and100-seed weight changed a little. The yield enhancementof varieties from Jilin-Liaoning province was the result of increased100-seed weight; and that ofvarieties from Yellow-Huai-Hai Rivers Valley was also associated with100-seed weight increament,while pod and seed numbers per plant decreased significantly. The traits of varieties fromJilin-Liaoning provinces changed the least compared with that in other regions.
2. Quality trait evolution of widely-planted varieties from different regions and decades
The contents of protein and isoflavones increased as the orignated latitudes of varieties weregetting lower. The content of oil, Vitamine E and carotenoids of vatieties from the northeast weremore than that of varieties from Yellow-Huai-Hai Rivers Valley. The contents of linoleic acid,linolenic adid, statch, oligosaccharide, isoflavone and lectin of varieties from the north part ofHeilongjiang province increased significantly with the released years; the contents of linoleic acid,palmatic acid, total isoflavones of varieties from the middle and south parts of Heilongjiang provinceincreased significantly along with the released decades; the starch content of varieties fromJilin-Liaoning provinces increased with the year of release, the contents of oleic and stearic adids, andlectin of varieties from Yellow-Huai-Hai Rivers Valley increased significantly with the released year.
3. Changes of photo-thermal response of widely-planted varieties derived from different regions and decades
The photoperiod sensitivity (PS) under spring and summer sowing of the varieties from the northpart of Heilongjiang province and Yellow-Huai-Hai Rivers Valley decreased with the decades ofrelease, while the same index of varieties from middle and south parts of Heilongjiang provincedeclined under summer sowing. The temperature sensitivity (TS) under short day of varieties from themiddle and south parts of Heilongjiang province and Yellow-Huai-Hai Rivers Valley decreased withthe released year, while that under long day of varieties from Yellow-Huai-Hai Rivers Valleyincreased with decades of release. The ΔPS(PS difference of under different temperature), ΔTS(TSdifference under different photoperiod) and PTCS (photothermal comprehensive sensitivity) ofvarieties from Yellow-Huai-Hai Rivers Valley decreased. The PS, TS and PTCS of widely-plantedvarieties increased gradually with the decreasing of their original latitude, while ΔPSand ΔTSattainedopposite trends compared with the former ones. It showed that the PS, TS and PTCS of varieties fromsouth are more sensitive than that from the north of China, but the photothermal interactive responsesof varieties were opposite.
4. The pedigree analysis of widely-planted varieties
Based on the collected pedigree data of the113soybean varieties, the end ancestors and directparents of the widely-planted varieties were traced. Compared to the data of the varieties reportedbefore, the number of end ancestors of widely-planted varieties was higher than the ordinary varieties,and their genetic bases were broader. Newly-bred widely-planted varieties have richer geneticbackground than the previous widely-planted varieties. Genetic basis of widely-planted soybeanvarieties from Yellow-Huai-Hai Rivers Valley are broader than that of ones from the northeast China.
5. The genetic diversity of widely-planted soybean varieties from different regions and decades
The average polymorphism information content of the varieties from the north part ofHeilongjiang province, south and middle parts of Heilongjiang province, Jilin-Liaoning province andYellow-Huai-Hai Rivers Valley were0.414,0.469,0.522and0.562, respectively. Except for thevarieties from the north part of Heilongjiang province, the varieties released after1980in the otherthree regions had higher PIC values than that of varieties released before1980. UPGMA-based clusteranalysis classified the tested varieties into three major groups, corresponding with the original placesof the varieties. The varieties from the northeast and Yellow-Huai-Hai Rivers Valley were classifiedinto two major groups respectively, except for Jidou12, a variety from the northern Yellow-Huai-HaiRivers Valley, which was classified into an independent group. In the northeast China, varieties ofHeilongjiang province were classified into one subgroup and the varieties from Jilin and Liaoningprovinces were into another subgroup, indicating that the widely-planted varieties from the sameregion were apparently homogenized.
6. The exploitation of elite alleles of varieties from different regions and decades
Among the SSR markers identified in this study, there were141loci associated with the24traits,39loci associated with the yield traits including yield, pods and seeds per plant and100-seed weight19loci associated with plant height, lodging index, the days to flowering and maturity, and82loci associated with the quality traits including protein, oil and isoflavones. The analysis on marker effectsand the identification of positive and negative loci and carriering materials showed that the extremeaccessions possessed the alleles with larger effects.
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