不同生态条件下籼粳交后代亚种分化研究及环境响应
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摘要
以利用不同亚种有利基因为目标的籼粳稻杂交是水稻重要育种方法,在迄今国内外对亚洲栽培稻分类、起源、演化以及籼粳稻杂交育种进行广泛研究并取得重大成果的基础上,本课题以典型籼粳稻杂交(七山占×秋光)重组自交系(F6)和分离世代(F2和F3)群体为试材,在不同生态环境下种植(华南籼稻区广东和东北粳稻区辽宁,F2分别采用混合法、一粒传法和系谱法处理),采用形态标记(程氏指数法)和分子标记(InDels和ILPs)相结合的方法,比较研究不同生态条件下形态分化、遗传分化、经济性状的差异,明确不同生态条件下籼粳稻杂交后代亚种分化规律及其与经济性状的关系,揭示自然选择与人工选择对亚种间杂交后代籼粳分化的影响及其遗传基础,探索利用亚种分化与经济性状的关系评估不同生态地区籼粳适应性的可能性与方法,为提高籼粳稻杂交育种效率提供科学依据,为实施“籼稻改粳稻”提供理论指导。
     1.在低世代群体中,各群体亚种属性都呈连续性变异且正态分布,并且两生态地区F2代群体间、两地区F3代单粒传法群体间,以及F2代和F3单粒传法群体间都无偏分离现象;而在两地区F3代混合法群体间、F3代单粒传法与混合法群体间,以及F2代与F3代混合法群间都出现显著的差异及明显的偏分现象。并且发现F3代群体程氏指数法与分子标记法所得的分类结果,无论是在辽宁和广东两个生态地区间还是混合法及单粒传法两个后代处理方法上,总体水平都是一致的,而且Dj与稃毛、1~2穗节长、籽粒长宽比及程氏指数上都呈极显著的相关性。
     2.在F6代群体中,不同地区及后代处理方法的群体同样呈连续性变异且正态分布,并且两地区单粒传法群体在分布上无差异(P=0.93);混合法群体在分布上有微小差异但不显著(P=0.40);系谱法群体呈极显著差异(P=0),且两群体明显偏向籼型,而广东地区群体比辽宁更为偏籼。并且,在混合法和单粒传法中,籼粳血缘Dj从偏籼或偏粳个体到中间血缘个体在频次分布上不呈连续性递增或递减,而是极大部分籼粳血缘集中于中间值(40%-60%),而偏籼或偏粳个体(小于40%或大于60%)骤然减少,该现象说明籼粳交育种存在综合亚种优势的潜力。
     3.在亚种属性变化的研究中发现,1-2穗节长在形态学水平与亚种分类关系密切,可作为形态学分类的有效指标,而壳色在分子水平上与亚种遗传分化关系密切,可作为籼粳遗传分化的形态学辅助指标,而籽粒长宽比在形态学及分子水平上都与亚种分化有紧密的关系,可作为籼粳交后代亚种分化中的一个有效指标,来定向性的指示其分化过程中亚种的变化。另外,我们还发现籽粒长宽比在辽宁和广东地区间有显著的差异,说明该性状在环境响应上反应迅速,推测其可能还受表观遗传影响。
     4.在研究籼粳亚种分化与经济性状的关系中,发现群体的千粒重与亚种属性有关,呈粳型>偏粳型>偏籼型>籼型的性状表现,且与千粒重有关的粒型及粒重基因区域血缘还与Dj呈显著的正相关;而结实率和单株产量却与生态适应性有关,因为其在相关性分析中和性状变化上呈现在辽宁地区粳型的性状表现较好,而在广东地区表现相反的现象;在F6代群体中,发现籼粳分化不仅与粒型及粒重基因有关,还与一些株型、穗型和抽穗期等性状基因有极显著相关。
     上述结果表明,遗传重组作用不会引起群体的偏分离,环境效应主要是影响群体中不同籼粳血缘成分在不同生态条件下的稳定或纯合速率,是生态适应性的体现,其选择及淘汰作用较弱,影响稳定群体偏分离的作用并不显著。而人工选择才是造成籼粳交后代偏分离的主要原因,并伴随着自然选择和生态适应性的影响,在群体稳定后产生显著的偏分离现象。另外,籼粳分化还与一些重要农艺性状有紧密的关系,尤其是在粒型及粒重相关性状上,无论是在性状表现上,还是在相关性上都呈现紧密的关系。这可能是籼粳交育种虽综合了亚种间血缘,但却并未达到综合亚种优势性状效果的直接原因。
Indica-japonica hybridization is an important rice breeding method, aimed at taking advantage of favorable genes in different subspecies. To date, great achievements have been obtained in the wide study on classification, origin, and develpment of Asian cultivated rice and indica-japonica hybridization breeding. Based on above, this study uses F2, F3, and F6generations population from crossing of typical indica QiShanZhan and japonica Akihikari as the materials, which are grown in Liaoning (japonica area in Northeast China) and Guangdong(indica area in South China) province, respectively. Begining at F2generation, populations are harvested by single-seed descent (SSD), bulk harvesting (BM), and pedigree methods (PM). Using morphological marker (Cheng's Index) and molecular markers (InDels and ILPs), a comparative study has been made on the differences between morphological and genetic differentiation, and economic traits in different environments. In addition, this study also identifies the law of subspecies differentiation in filial generation of indica-japonica hybridization, and its relationships with economic traits under different environments; reveal the genetic basis on the effect of natural and artificial selection on subspecies differentiation; explores how to estimate indica-japonica adaptability of different environments, using the relationship between subspecies differentiation and economic traits; and finally, provides new insights into the process of subspecies differentiation to improve the efficiency of indica-japonica hybridization breeding, and inform to carry out" changing indica to japonica" project.
     1. In early generations, subspecies characteristics of all population show the normal distribution and continuous variation, and no distortion segregation is found between two environments of F2population and F3SSD populations, and F2and F3SSD population. However, there are significant differences and obvious phenomenon of distortion segregation between two environments of F3BM population, SSD and BM population of F3generation, and F2and F3BM population. In addition, it is found that the results classified by Cheng's Index are in accordance with those classified by molecular markers, whether in different environments or seed harvesting methods. Furthermore, there are also significant correlations with distribution of japonica kinship percentage (Dj) found in glume pubescence (GP), length of the first and second panicle internodes (LPI), length-width ratio of grains (LWR), and Cheng's Index.
     2. In F6generation, populations also show a normal distribution and continuous variation. According to comparing the population distribution of different environments, it is found that there's no difference in SSD populations (p=0.93), a minute but non-significant difference in BM populations (P=0.40), a significant difference in PM populations (P=0). And the whole PM populations are indica-deviated; moreover, PM population in Guangdong is more indica-deviated than that in Liaoning. Besides, in BM and SSD population, there's a discontinuous variation from indica-deviated or japobica-deviated individuals to neutral kinship individuals in frequency distribution; the most majority of individuals are neutral kinship (Dj=40%-60%), but few is indica-deviated or japonica-deviated (Dj>60%or<40%). This phenomenon suggests that there are potentials existing in indica-japonica hybridization breeding.
     3. Drawing on the study on changes of subspecies characteristics, LPI is found to be closely related with morphological subspecies differentiation, probably as a effective index of morphological classification; but glume color (GP) has a close relationship with subspecies differentiation in molecular level, probably as an accessory index in morphology for subspecies genetic differentiation; additionally LWR is closely related with subspecies differentiation both in morphological and molecular level, possibly as an effective classification index to indicate the changes of subspecies in the process of differentiation. In addition, LWR also shows great differences between Liaoning and Guangdong provinces, demonstrated to has a fast response to environment, which implies that LWR is probably affected by epigenetics.
     4. According to the study on the relationships between subspecies differentiation and economic traits,1000-grain weight (KW) is related to subspecies characteristics and shows japonica type>japonica-deviated type> indica-deviated type> indica type; and the kinship in the region of grain weight and shape genes with respect to KW is significantly and positively correlated with Dj; but seed setting rate (SSR) and grain yield per plant (GYP) are related with ecological adaptability, because correlation results show that their performance of japonica type is better than indica type in Liaoning but it is opposite in Guangdong; in F6generation, not only grain shape and weight genes are related to subspecies differentiation but some other genes with respect to plant type, panicle type, and heading date et al. are also significantly correlated with subspecies differentiation.
     The results above demonstrate that genetic recombination could not cause distortion segregation; and environment could affect the speed of stabilization on different kinship individuals, which is the performance of ecological adaptability; and in the process of population differentiation, selection and elimination effect of environment are weak, which could not significantly affect population differentiation and cause distortion segregation. However, artificial selection is found as the main factor obviously causing distortion segregation in filial generation from indica-japonica hybridization, with the effect of nature selection and ecological adaptability. In addition, it is also found that there's close relationship between subspecies differentiation and some important agronomic traits. Especially in grain shape and weight, they show a close relationship whether in traits performance or correlation. This may be the primary cause resulting in why indica-japonica hybridization breeding didn't achieve the required effect of combining subspecies superiority but subspecies kinship.
引文
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