本文關(guān)鍵詞：高粱馴化相關(guān)性狀遺傳結(jié)構(gòu)的解析　出處：《中國農(nóng)業(yè)大學(xué)》2016年博士論文　論文類型：學(xué)位論文

  更多相關(guān)文章： 馴化 高梁 QTL 抽穗期 株高

【摘要】：作物的馴化始于12000年前的新石器革命,在此以后陸續(xù)有多種作物在不同地區(qū)由不同的古人類完成了馴化。作為世界五大作物之一,高粱(Sorghum bicolor (L) Moench)起源于非洲東北部的蘇丹、埃塞俄比亞及其周邊地區(qū)。在其馴化的過程中,高粱發(fā)生了一系列形態(tài)學(xué)和生理學(xué)巨大變化,這些變化包括落粒性的喪失、穎殼顏色和種皮顏色變化、開花期和成熟期的同步、籽粒增大以及株型更加緊湊等等。在本研究中,我們利用野生高粱SV (Sorghum virgatum)×栽培高粱Tx623組配的F2群體和F2：3群體,對高粱馴化相關(guān)性狀進行了QTL定位,分別對位于10號染色體上的抽穗期主效位點以及位于9號和1號染色體上兩個株高主效位點進行了精細(xì)定位,并對候選基因在高粱馴化中的作用進行了解析。本文主要研究結(jié)果如下：1.利用R/qtl軟件包及272個F2單株,構(gòu)建了一套包含均勻分布于高粱10條染色體上的216個SSR標(biāo)記的分子遺傳圖譜。圖譜總長1610.5 cM,相鄰兩個標(biāo)記的平均遺傳距離是7.8 cM。2.對F2群體(北京)的18個馴化相關(guān)性狀及F2：3群體(海南)的6個性狀進行表型調(diào)查,利用R/qtl軟件包,對這些性狀進行大規(guī)模的QTL定位,共檢測到66個相關(guān)QTLs。這些位點分布于高粱的10條染色體上,但其分布并不均勻,在某些染色體區(qū)段上“富集”。對QTL加性效應(yīng)分析表明,來自栽培種親本的正向效應(yīng)多有利于葉長、葉寬及粒重、粒寬增加等,而來自野生種親本的正向效應(yīng)多有利于株高、分蘗的增加等。3.在前期F2群體及F2；3群體初定位的基礎(chǔ)上,我們對位于10號染色體上控制抽穗期的主效QTL-HD1進行精細(xì)定位,利用重組個體后代驗證的方法將定位區(qū)間縮小到37 kb的物理區(qū)間內(nèi)。在此區(qū)間內(nèi),編碼CONSTANS-like蛋白的Sb10g010050對應(yīng)于HDI基因,野生親本SV的HDI基因的第一個外顯子上存在一個5-bp的缺失,導(dǎo)致了HD1蛋白缺失了鋅指結(jié)構(gòu)和CCT結(jié)構(gòu)功能域,從而引起蛋白功能的喪失。DNA多態(tài)性分析發(fā)現(xiàn)栽培種的多態(tài)性明顯低于野生種,Tajima'sD檢測結(jié)果表明栽培種該基因5-bp缺失位點受到輕微平衡選擇(Tajima D's test=2.39, P 0.05)。比較基因組分析發(fā)現(xiàn),在高粱HDl位點的對應(yīng)于玉米的9號染色體、水稻的6號染色體及谷子的4號染色體區(qū)段,且在水稻6號染色體及谷子的4號染色體相關(guān)片段上都定位到了抽穗期相關(guān)QTL。對谷子的HD1基因的大規(guī)模測序發(fā)現(xiàn),一個位于剪切位點的變異(GT→AT)較為常見(78/98),造成剪切位點從787bp的“GT”前移到754bp處的“GT"。關(guān)聯(lián)分析在剪切位點處檢測到顯著關(guān)聯(lián)信號(P=9.50×104)。谷子HD1基因的核酸多樣性分析發(fā)現(xiàn)栽培谷子的DNA多樣性顯著降低。這些結(jié)果表明HD1基因在高粱、水稻和谷子中受到平行選擇。4.根據(jù)F2和F2：3初定位結(jié)果,在9號染色體上檢測到控制株高及其他性狀的主效QTL(dwl)。利用連續(xù)重組后代驗證的方法對其進行精細(xì)定位,將定位區(qū)間縮小到13.98 kb的物理區(qū)間內(nèi)。對該區(qū)間內(nèi)的兩親本的基因進行測序,發(fā)現(xiàn)栽培親本Tx623的基因Sobic.009G229800第二個外顯子上存在A→T的SNP變異,導(dǎo)致翻譯提前終止。對不同來源的高粱品種候選基因測序分析發(fā)現(xiàn)在來自美國、印度等地的矮化栽培高粱中提前終止變異時很常見的,且這些矮化品種的DW1基因核酸序列間不存在任何變異,多樣性低,說明這個A到T的突變發(fā)生在近期。然而,不存在功能位點改變的栽培高粱與野生高粱相比,DNA多樣性并不存在顯著降低,表明dwl基因在馴化過程中可能并未受到選擇,功能位點突變可能是現(xiàn)代育種的結(jié)果。5.對1號染色體上的株高QTL-ph1進行精細(xì)定位,將定位區(qū)間縮小到24.5-kb的定位區(qū)間內(nèi),區(qū)間內(nèi)僅存在一個基因,Sobic.001G106000(kn2),該基因編碼一個KNOTTED-like轉(zhuǎn)錄因子。親本DNA測序發(fā)現(xiàn)兩個能引起編碼氨基酸變化的SNP變異,但其可能并非是導(dǎo)致表型變異的相關(guān)功能位點。
[Abstract]:The domestication of crops began in the Neolithic Revolution 12000 years ago, after which a variety of crops have been domesticated by different kinds of ancient people in different regions. As one of the five largest crops in the world, sorghum (Sorghum bicolor (L) Moench) originated in Sultan, Ethiopia and its surrounding areas in northeastern Africa. During its domestication, sorghum has undergone a series of dramatic changes in morphology and physiology. These changes include loss of grain, the color of shell and seed coat, the synchronization of flowering and maturity, the increase of grain and the more compact plant type. In this study, we use wild sorghum SV (Sorghum virgatum) * Tx623 group with cultivation sorghum F2 population and F2:3 population, the sorghum domestication related traits were QTL, are located in heading period of major loci on chromosome 10 and chromosome 1 and 9 in two main plant height the fine mapping of loci, and the role of candidate genes in Sorghum domestication were analyzed. The main results of this study are as follows: 1.. Using R/qtl software package and 272 F2 individuals, we constructed a set of molecular genetic map containing 216 SSR markers distributed evenly on 10 chromosomes of sorghum. The total length of the map was 1610.5 cM, and the average genetic distance of two adjacent markers was 7.8 cM. 2. phenotypic investigation of 18 domesticated traits and 6 traits of F2:3 population (Hainan) in F2 group (Beijing). A large scale QTL localization of these traits was carried out by R/qtl software package, and 66 related QTLs were detected. These loci are distributed on 10 chromosomes of sorghum, but their distribution is not uniform and "enriched" on some chromosomal segments. The analysis of additive effect of QTL showed that the positive effects of parents from cultivars were more conducive to leaf length, Ye Kuan and grain weight and grain width. 3., on the basis of early F2 population and F2 and 3 population initial location, we fine positioned the dominant QTL-HD1 on chromosome 10, and reduced the location interval to 37 KB by using the method of recombinant offspring. In this interval, the Sb10g010050 encoding CONSTANS-like protein corresponds to HDI gene. There is a deletion of 5-bp on the first exon of HDI gene of wild parent SV, which results in the loss of zinc finger structure and CCT functional domain, which results in the loss of protein function. DNA polymorphism analysis showed that the polymorphism of cultivated species was significantly lower than that of wild species. Tajima'sD detection showed that the 5-bp deletion site of cultivated species was slightly balanced (Tajima D's test=2.39, P 0.05). Comparative genomic analysis revealed that there was a heading related QTL in maize sorghum HDl locus corresponding to maize chromosome 9, rice chromosome 6 and millet chromosome 4, and on chromosome 4 of rice chromosome 6 and millet. A large-scale sequencing of HD1 gene in Millet showed that a mutation located at the shear site (GT to AT) was more common (78/98), causing the shear site to move from 787bp GT to 754bp at 754bp. A significant correlation signal (P=9.50 x 104) was detected at the shear site by correlation analysis. The analysis of nucleic acid diversity of the HD1 gene of millet showed that the DNA diversity of cultivated millet was significantly reduced. These results showed that the HD1 gene was selected in sorghum, rice and millet. 4. according to the initial results of F2 and F2:3, the main effect QTL (DWL) of control plant height and other traits were detected on chromosome 9. The positioning interval is narrowed to the 13.98 KB physical interval by using the method of continuous recombined verification. According to the sequencing of the two parents in the interval, it was found that there existed SNP variation of A to T on the second exons of the parental gene Tx623 Sobic.009G229800, leading to the early termination of the translation. The sorghum candidate gene sequencing analysis found in different sources from the United States, India and other places of dwarfing cultivation sorghum early termination mutation is common, and these dwarf varieties of DW1 gene sequence, no mutation, low diversity, the A to T mutation in the near future. However, there was no significant decrease in DNA diversity in cultivated sorghum compared with wild sorghum without functional site change, indicating that DWL gene may not be selected during domestication. Functional site mutation may be the result of modern breeding. 5. fine mapping of plant height QTL-ph1 on chromosome 1, narrowing the location interval to the 24.5-kb location interval. There is only one gene, Sobic.001G106000 (kn2), which encodes a KNOTTED-like transcription factor. Parent DNA sequencing found two SNP mutations that could cause the changes in encoded amino acids, but may not be related functional sites that lead to phenotypic variation.
【學(xué)位授予單位】：中國農(nóng)業(yè)大學(xué)
【學(xué)位級別】：博士
【學(xué)位授予年份】：2016
【分類號】：S514

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