摘要
作物种质资源具有广泛的遗传变异,是选育突破性新品种的物质基础,育成品种是经多年育种与生产实践证明最为重要的种质资源。我国1923-2005年共育成了1300个大豆育成品种,研究这些育成品种的遗传基础及其近年来的演变趋势,对指导中国大豆品种改良具有重要的现实意义。
系谱分析能较好地阐明作物育成品种的整体遗传基础,并且具有经济、简便等优点；我国近百年大豆育种实践资料蕴涵丰富的系谱、亲本选配等信息,是研究我国大豆育成品种整体遗传基础的理想材料。本研究利用中国大豆育种系谱资料分析中国大豆育成品种的整体遗传基础,对10个主要家族的育成品种,利用均匀分布于大豆20个连锁群的161对简单重复序列(SSR)标记引物进行SSR分子标记研究,综合亲本系数(CP)分析育成品种间的遗传相似性和亲缘关系,揭示其遗传基础,以期为我国大豆育种亲本选配,扩大亲本来源以至拓宽我国大豆育成品种遗传基础提供参考。主要结果如下：
1.我国1923-2005年所育成1300个大豆品种中,北方一熟春豆生态区(Ⅰ区)、黄淮海二熟春夏豆生态区(Ⅱ区)、长江中下游二熟春夏豆生态区(Ⅲ区)、中南多熟春夏秋豆生态区(Ⅳ区)、西南高原二熟春夏豆生态区(Ⅴ区)和华南热带多熟四季大豆生态区(Ⅵ区)分别有682、395、64、120、16和23个,其中1019、202、70和9个分别由杂交、系统选择、诱变和转DNA育种方法育成。1300个品种的细胞核和细胞质基因库分别追溯到670和344个祖先亲本,一些重要祖先亲本育成了早期骨干品种(如满仓金、紫花四号、徐豆1号、齐黄1号和南农493-1等),这些品种又育成新的品种,经过多轮育种过程形成复杂的祖先亲本家族,一些重要祖先亲本在Ⅰ区已经历5-8轮的育种过程,Ⅱ区也经历了5-7轮的育种过程,Ⅲ、Ⅳ、Ⅴ、Ⅵ区经历了3-5轮的育种过程。
我国大豆杂交育成品种共使用1391个直接亲本,绝大多数为育成品种和育种品系,少量为地方品种与外国品种。生产上常采用性状优良适合当地条件的育成品种作为供体亲本以加速育种进程,如：合丰25、吉林20、徐豆1号、齐黄1号、矮脚早、南农493-1和十胜长叶等作为直接或间接亲本多次使用,形成一些重要亲本。我国大豆杂交育种的组配方式的演变发展趋向于以育成品种(C)和育种品系(S)相互组合(CxC、 SxS、c×s、SxC)的四种组配为主。全国71.78%的杂交育成品种是以这四种组配方式育成,近十年这四种组配方式更高,为83.50%。
2.670个祖先亲本源自我国六个生态区及国外,其中Ⅰ、Ⅱ、Ⅲ生态区和国外的祖先亲本提供了主要核质遗传贡献,各生态区间的种质交流较少,以本生态区的祖先亲本的遗传基础为主,其它生态区的祖先亲本遗传贡献较少。大部分祖先亲本只衍生1-2个育成品种,少数重要祖先亲本,如：Ⅰ区的金元(A146)、四粒黄(A210)和白眉(A019),Ⅱ区的滨海大白花(A034)、山东张寿地方品种(A295)、滑县大绿豆(A122)、铜山天鹅蛋(A231)、即墨油豆(A133),Ⅲ区的奉贤穗稻黄(A084)、51-83(A002)、上海六月白(A201)等祖先亲本经多轮育种过程衍生出大量品种,形成了庞大祖先亲本家族。综合衍生品种数、细胞核、质遗传贡献值和衍生品种轮数四项指标,从670个祖先亲本中筛选出113个对中国大豆育成品种核心祖先亲本,这些祖先亲本来自国内六生态区和国外。113份祖先亲本(占总数的16.87%)对1300个育成品种的核、质遗传贡献分别占70.90%和74.85%；这些核心祖先亲本对大豆核心种质收集、利用以及亲本选配具有重要参考意义。
3.各生态区育成品种的平均每个育成品种使用祖先亲本数在逐渐增加,其中Ⅰ、Ⅱ区尤其明显,六生态区1996-2005年育成品种的平均每个育成品种使用祖先亲本数较1986-1995年的增加近一倍,近期育成品种比早期育成品种的遗传基础较为宽广,这与大量育成品种(品系)作为杂交育种亲本有关。经过10年的育种进程我国大豆育成品种祖先亲本扩大近一倍,地理来源更广泛,各生态区育成品种的国外和异生态区祖先亲本的核质遗传贡献有所增加,整体上我国大豆育成品种的遗传基础有所拓宽。但单个祖先亲本对我国大豆育成品种的遗传贡献不均衡现象有所加剧,遗传贡献向一些重要祖先亲本集中。如：Ⅰ区的A146、A210和A019,Ⅱ区的A034、A231、A133,Ⅱ区的A084、A002、A201等少数祖先亲本衍生品种数较1923-1995间增加一倍多,遗传贡献值增加一半以上。
4.亲本系数分析表明近期育成品种(1986-2005)较早期育成品种(1923-1985)遗传基础更为宽广,受试品种中Ⅱ区的品种遗传基础最为宽广,Ⅲ区的品种遗传基础较狭窄。各家族育成品种中,Ⅱ区的A295、A122、A231和A133四个家族系谱复杂,涉及祖先亲本多,遗传基础较为宽广。Ⅲ区一些家族如A084、A002、A201和矮脚早(A291)由于选择育种较多,亲本系数平均值较大,遗传基础较为狭窄。
5.基于SSR标记的遗传多样性、互补等位变异和特异等位变异研究表明：近期品种(1986～2005)较早期品种(1923～1985)遗传多态性信息含量(PIC)丰富,平均和特有等位点多、特缺等位点少,遗传互补等位变异点差异大,两者遗传关系最远,近期品种较早期品种遗传基础更为宽广。育成品种间的遗传基础与其所在的地理区域分布有关,Ⅰ和Ⅳ生态区地理位置相隔较远,互补等位变异点数最多,遗传关系也最远,Ⅱ、Ⅲ两生态区地理位置较近,种质交流较多,互补等位变异点数最少,遗传关系最近。
各家族育成品种的SSR标记的遗传多样性、群体遗传结构、遗传相似性和特异等位变异研究表明各家族品种间的遗传基础与其所在的地理分布有关。A201和A291家族的品种主要分布于Ⅲ区的湖南、湖北和四川等地遗传背景较为一致；A002和A084家族的品种主要分布于江苏、河南和四川等地其遗传背景也较为一致；Ⅱ区的A034和A231家族的品种主要分布于江苏、河南和四川等地其遗传背景较为一致,A133和A295家族的品种主要分布于山东和河南等地其遗传背景较为一致。东北白眉(A019)家族和其它9个家族地理距离较远,存在较多特有、特缺等位点,而Ⅲ区和Ⅱ区地理位置较近,种质交流较多,两区祖先亲本家族间特有、特缺等位点数较少。其中Ⅲ区A002和Ⅱ区的A231和A122间无特有等位点,Ⅲ区A084、A201和Ⅱ区的A034、A231亚群间无特缺等位变异点。
6.亲本系数是静态地假定双亲遗传贡献相同,而SSR标记是动态地反映品种间DNA水平的遗传差异。10个家族的亲本系数和SSR相似性系数矩阵Mantel检验相关验证表明两者相关程度较低。通过对37个杂交组合的SSR标记数据分析证明基于SSR标记的遗传贡献率法(GC)与相似系数法(SC)两种遗传贡献计算方法结果十分接近,母本对子代的遗传都明显高于父本。两法和基于系谱资料计算遗传贡献率的亲本系数法(CP)的计算结果差别较大。对37个杂交组合的等位变异点的传递与遗传重组分析表明：母本传递给子代的位点数多于父本,20个连锁群中C2、A2、J、F连锁群传递位点数较多, Ⅰ连锁群最少；母本重组发生率明显高于父本,近期品种重组率高于早期品种。
Crop germplasm with abundant genetic variant is basic material for crop breeding program. The long-term breeding practice fully prove that crop released cultivars were important germplasm. Total1300soybean cultivars had been released in China from1923to2005. The knowledge of genetic base and their change for recent years of soybean cultivars can be useful for setting the approaches and steps of genetic base-broadening in soybeans in China.
The genetic base of crop was elucidated efficient by the pedigree analysis, especially the population genetic base of crop; The soybean breeding pedigree data with abundant information of pedigree and parents selection for approximate hundred years was ideal for the their population genetic base. Based on an analysis of pedigree data of the1300soybean cultivars released, simple sequence repeats (SSR) molecular marker161pairs of primers located on well-distributed20molecular linkage groups (MLG) and coefficient of parents (CP). The objective of this study is to make a thorough analysis on the population genetic bases of soybean released cultivars; summarize the relation between geographical source and genetic contribution of ancestor from soybean cultivars released in various decade; determine genetic similarity and relationship among cultivars; so as to provide relevant information for further breeding plans of soybeans parents selection and enlarge sources parents up to broadening the genetic base released cultivar. The main results were as follows:
1. Total1300soybean cultivars were released in China from1923to2005,682,395,64,120,16and23were released in Northern China single cropping spring planting varietal eco-region (I), Huang-Huai-Hai double cropping spring and summer planting varietal eco-region (II), Middle and Lower Changjiang valleys double cropping spring and summer planting varietal eco-region (III), Central-south multiple cropping spring summer and fall planting varietal eco-region(IV), Southwest plateau spring and summer planting varietal eco-region (V) and South China tropics multiple cropping all season planting eco-region (VI),1019、202、70and9were developed from artificial hybridization, natural variant selection, mutation and transfer DNA, respectively. These cultivars was traced back to670nuclear and344cytoplasmic ancestors; Some major ancestors were cultivated early major cultivars (such as Mancangjin, Zihua si hao, Xudou1hao, Qihuang1hao and Nannong493-1), which were bred more new cultivars, after many breeding cycles, major ancestors were developed the complicated ancestor family. Nowadays some major ancestor from I, II and III, IV, V, VI eco-region go through5-8,5-7and3-5breeding cycles, respectively.
1391soybean accessions were selected with direct crossing parent, which included the majority of cultivars and breeding lines and a little landraces and foreign countries cultivar. In order to process of breeding, some important cultivars (such as Hefeng25、Jilin20, Xudou1hao, Qihuang1hao, Aijiaozao, Nannong493-land Shishengchangye etc.) were used for direct crossing parents time after time. Parental combination of71.78%soybean cultivars released were the four parent combination between cultivars (C) with breeding line(S)(CxC、SxS、C×S、SxC) in China, and were more83.5%recent ten years.
2. Total670ancestors were composed of landraces, breeding lines, improved cultivars, wild accessions and uncertain type, come from Eco-region I, Ⅱ, Ⅲ,IV, V, V I and foreign countries. Both nuclear and cytoplasmic germplasm from Eco-region I, II, III and foreign countries accounted for most part of the ancestry of the released cultivars. The exchange and utilization of germplasm among eco-regions were poor and mostly still limited in their own eco-regions. The most part of the ancestors was used in only1-2released cultivars, while only a few ancestors provided large genetic contribution but still limited basically in their local ancestors. Such as from eco-regions I Jinyuan(A146), Silihuang (A196), Baimei (A019), Ⅱ Binhaidabaihua(A034), Shandongzhangshou landrace (A295), Huaxiandaludou (A122), Tongshantianedan(A231), Jimoyoudou(A133), III (Fengxiansuidaohuang(A084),51-83(A002), Shanghailiuyuebai(A201). The113core ancestors out of the670were nominated according to the number of derived cultivars, nuclear genetic contribution, cytoplasmic genetic contribution and number of breeding cycles of each ancestor. Among them,34,28,10,14,4,3and20were from I, Ⅱ, Ⅲ, IV, V, VI eco-region and abroad, respectively. It accounted for16.87%of the total ancestors and provided70.90%nuclear and74.85%cytoplasmic genetic contribution to the whole released1300cultivars.
3. The average number of ancestors per cultivar released between1996to2005in various eco-regions was about double or more of that between1986to1995. The genetic base of a single cultivar is relatively narrow for most of the released cultivars but has been gradually broadened, especially in Eco-region Ⅰ and Ⅱ; and the average number of ancestors per cultivar released between1996and2005in various eco-regions was approximately double of the period between1986and1995. The of pedigree of recent cultivars were more complication than early ones because scale of cross-breeding and parent combination between cultivars with breeding line raise recent years.
4. The analysis of parentage coefficient showed that the genetic base of recent cultivars (1923-1985) were more broad than early ones. The genetic base of cultivars for Ⅱ eco-region was most broad, and Ⅲ eco-region was most narrow among Eco-region Ⅰ, Ⅱ, Ⅲ and Ⅳ. Among ten ancestors family, the genetic bases of Ⅱ eco-region such as A295, A122, A231and133family released cultivars is broad, Ⅲ eco-region such as A084, A002, A201and Aijiaozao (A291)family were narrow.
5. Based on analysis of SSR marker of polymorphism information content (PIC) and alleles, we find that The specifically existent alleles, complementary alleles and PIC of recent cultivars (1986～2005) was more than early cultivars (1923～1975), and the specifically deficient alleles was low. The results showed that the genetic base of recent cultivars (1923-1985) were more broad than early ones. The genetic base of cultivars were was related to geographical district of cultivars. The geographical district Ⅰ and Ⅱ eco-region is far and the number of complementary alleles were most, so the genetic relationship between them and was far. On the contrary,The geographical district Ⅰ and Ⅱ eco-region is near and exchange of germplasm was much, the number of complementary alleles were low, so the genetic relationship between them and was close.
The result of analysis of genetic structure of the population, specifically existent alleles and genetic similarity were indicated that the genetic base of cultivars were was related to geographical district of cultivars. For example, A201and A291ancestor families cultivars from south region was the geographic distribution in Hunan, Hubei, Sichuan and Jiangxi province and was almost clustered in same group.
The number of alleles of A201and A291ancestor family from south regions was more low than ones of A034, A231and A133ancestor family from Ⅱ region. Genetic diversity level in recent cultivars (1986～2005) was higher than that in early cultivars (1923～1975). The different geographical district cultivars can be differentiated by two methods and reveal the distribution of ancestor family characteristic. For example, A201and A291ancestor families cultivars from south region was the geographic distribution in Hunan, Hubei, Sichuan and Jiangxi province and was almost clustered in same group.
6. The average genetic similarity indicated the genetic variant and relationships of among cultivars in DNA level. The degree of relationship between the similarity matrices based on SSR and pedigree was measured by comparing the similarity matrices with the normalized Mantel test. A low positive correlation between the two matrices was observed. based on pedigree analysis and genetic similarity SSR markers was different. Parental genetic contribution can be estimated by from pedigree or from molecular marker data; there are three ways to calculate CP by pedigree, GC and SC by marker. We studied37soybean inbreeds with known pedigrees. The inbreeds were genotyped using161SSR markers. Parental contributions were estimated from marker similarity among an inbred and both of its parents, and were subsequently used to estimate CP. Estimates of parental contribution differed significantly between pedigree data and marker. The parental genetic contribution estimated from GC and SC was highly correlated. We concluded that female parental contribution estimated from marker data was more than male one when both parents were same one from pedigree data.
The study of distribution of recombination events show that the number of alleles from female parent were more than male parent. There was the difference of number of transition alleles among linkage groups, the linkage group C2, A2, J and F have much more number of transition alleles, but I was least. Recombination events of latest cultivars was more than old one.
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