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基于連鎖和關(guān)聯(lián)分析對水稻柱頭外露率的研究

發(fā)布時間:2018-07-24 13:51
【摘要】:水稻作為發(fā)展中國家最主要的糧食作物之一,提供著一半以上世界人口的主食。隨著人口的飛速增長,提高水稻產(chǎn)量的意義也變得愈發(fā)重要,而雜交水稻的推廣是提高水稻產(chǎn)量的一個可行辦法。雖然雜交水稻已經(jīng)大范圍商業(yè)化,但是不育系的低異交率使得雜交水稻的制種效率普遍偏低和制種成本的增加,從而限制了雜交水稻的推廣。柱頭外露是影響水稻異交性能的重要因子,柱頭外露的穎花的結(jié)實率是柱頭不外露的穎花的三倍以上,因此改良水稻不育系的柱頭外露率可以提高其異交率。近年來學(xué)者們定位了許多柱頭外露的QTL,但是大部分效應(yīng)都非常的小,目前也沒有利用柱頭外露QTL改良不育系柱頭外露的相關(guān)報道。在本研究中,我們通過連鎖分析定位了兩個大效應(yīng)的柱頭外露率QTL:q SE2和q SE6,通過關(guān)聯(lián)分析鑒定了三個柱頭外露QTL與三個粒形基因共定位,并驗證了它們的功能。本研究主要結(jié)果如下:1.通過構(gòu)建高柱頭外露率三系不育系超泰A和兩系不育系廣占63S的BC1F2群體,我們在第二染色體上定位到一個主效的柱頭外露QTL基因q SE2,解釋群體遺傳變異46%。NIL(超泰A)比NIL(廣占63S)的柱頭外露率高50%,且NIL(廣占63S)表現(xiàn)為花粉敗育,因此推斷q SE2就是光溫敏核不育基因tms5。2.利用溫敏不育基因tms5緊密連鎖的分子標(biāo)記鑒定Pokhreli和HD9802S的F2群體的育性,我們從中篩選出純合可育的192個單株用來構(gòu)建遺傳連鎖圖。在該群體中掃描到三個柱頭外露率的QTL,分別是q SE4、q SE6和q SE9。其中q SE6為主效的QTL,在群體中解釋柱頭外露率遺傳變異29%,但是該QTL來源于低值親本HD9802S。NIL(Pokhreli)比NIL(HD9802S)的柱頭外露率低30%。3.對529份多樣性栽培稻的花器性狀進行全基因組關(guān)聯(lián)分析,對柱頭外露相關(guān)性狀檢測到超過50個顯著性位點。秈稻和粳稻中檢測到的位點大多不同表明秈粳間柱頭外露率的遺傳差異較大。對于純系而言,柱頭外露率的遺傳力較高,屬于典型的數(shù)量性狀,受多個位點效應(yīng)的影響。應(yīng)用混合線性回歸,5個位點聯(lián)合解釋遺傳變異39%,表明柱頭外露存在很強的加性效應(yīng)。4.GWAS檢測到的三個關(guān)聯(lián)位點和三個粒形基因共定位,分別為GW2、GS3和GW5。利用這三個基因的三組轉(zhuǎn)基因材料,我們驗證了三個位點對柱頭外露的效應(yīng)。通過進一步考察三組轉(zhuǎn)基因材料的柱頭性狀,我們發(fā)現(xiàn)GS3會減小柱托長度,GW5會增加柱頭的大小,GW2會降低柱頭大小。三個基因?qū)λ玖P蔚挠绊懪c前人報道相符。因此三個基因通過調(diào)控穎殼和柱頭的形狀來影響柱頭外露。5.對529份栽培稻粒形的GWAS表明GS3和GW5是栽培稻粒形變異的主要基因。利用功能性標(biāo)記檢測529份材料的GS3和GW5的基因型,通過回歸分析我們發(fā)現(xiàn)兩個基因都能夠解釋柱頭外露遺傳變異20%以上,兩個基因聯(lián)合能夠解釋30%以上。這些結(jié)果表明,GS3和GW5也是控制栽培稻柱頭外露率的主要基因。6.為了研究GS3和GW5對于改良柱頭外露的可行性,我們將這兩個基因劃分為四種基因型組合:GW5GS3、GW5gs3、gw5GS3和gw5gs3。通過比較四種基因型組合的柱頭外露和產(chǎn)量性狀,我們發(fā)現(xiàn)GW5gs3這種基因組合具有最高的柱頭外露率和產(chǎn)量。而栽培稻中出現(xiàn)頻率最高的基因型組合gw5GS3的柱頭外露率和產(chǎn)量都最低,因此可以利用優(yōu)勢基因型組合GW5gs3對栽培稻進行遺傳改良。7.GW5gs3主要存在于秈稻II和熱帶粳稻中,而gw5GS3主要存在于秈稻I和溫帶粳稻中,這種分布的差異表明這GS3和GW5可能受到某種選擇。利用這兩個基因序列內(nèi)SNPs分析這兩個基因的連鎖不平衡,我們發(fā)現(xiàn)無論是否考慮群體結(jié)構(gòu),兩個基因間都存在較強的連鎖不平衡。通過對兩個基因不同基因型和不同基因型組合的核酸多態(tài)性分析,我們發(fā)現(xiàn)這兩個基因可能在育種的過程中受到選擇,從而形成不同亞群中GW5gs3和gw5GS3分布的差異,進而導(dǎo)致不同亞群的柱頭外露率和產(chǎn)量的差異。
[Abstract]:As one of the most important food crops in developing countries, rice provides more than half of the staple food of the world population. With the rapid growth of the population, the significance of increasing rice yield is becoming more and more important, and the popularization of hybrid rice is a feasible way to improve rice yield. The low cross rate of the breeding lines makes the seed production efficiency of hybrid rice generally low and the cost of seed production increasing, which restricts the popularization of hybrid rice. The exposure of stigma is an important factor affecting the cross cross performance of rice. The seed setting rate of the exposed spikelet is more than three times that of the unexposed spikelet, thus improving the stigma outside the sterile line of rice. The exposure rate can improve the rate of incrossing. In recent years, scholars have located many QTL of stigma exserted, but most of the effects are very small. There is no report on the use of stigma exsertion QTL to improve the exposure of stigma in sterility lines. In this study, we identified two major effects of the exposure rate of stigma QTL:q SE2 and Q SE6 by linkage analysis. The co localization of three stigma exserted QTL and three grain shape genes was identified and their functions were verified by correlation analysis. The main results of this study were as follows: 1. by building high stigma exposure rate three line sterile line super Thai A and two line sterile lines to occupy 63S BC1F2 population, we locate a main effective stigma exposed QTL on second chromophore. Gene Q SE2, which explains the population genetic variation 46%.NIL (A) is 50% higher than that of NIL (63S), and NIL (63S) shows pollen abortion. Therefore, it is concluded that Q SE2 is the fertility of the photoperiod sensitive genic sterility gene tms5.2. using thermo sensitive genic male sterile gene. 192 homozygous plants were screened to construct a genetic linkage map. In this group, the QTL of three stigma exposure rates was scanned, which were Q SE4, Q SE6 and Q SE9., which were the dominant Q SE6, which explained the genetic variation of the exposure rate of stigma 29% in the population, but the QTL was derived from the stigma of the low parent HD9802S.NIL. The total genomic correlation analysis was carried out on the flower characteristics of 529 varieties of cultivated rice with low exposure rate, and more than 50 significant loci were detected for the correlation of the stigma exposure. Most of the detected loci in Indica and japonica showed that the genetic difference between the exposure rate of the stigma between Indica and japonica was larger. For pure lines, the inheritance of the exposure rate of the stigma. High force, which belongs to the typical quantitative trait, is influenced by multiple locus effects. Mixed linear regression and 5 loci combined to explain genetic variation 39%, it indicates that the stigma exsertion has a strong additive effect.4.GWAS detected by three associated loci and three granular genes, which are GW2, GS3 and GW5. using three groups of these three genes, respectively. We verified the effect of three loci on stigma exposure. By further examining the stigma traits of three groups of genetically modified materials, we found that GS3 would reduce the length of the column, GW5 would increase the size of the stigma, and GW2 would reduce the size of the stigma. The effect of three genes on the grain shape of rice was consistent with the previous reports. Therefore three genes passed through the previous reports. The shape of the spikelet and the stigma was regulated to influence the GWAS of the grain shape of 529 cultivated rice with the stigma exsertion.5. indicating that GS3 and GW5 were the main genes of grain variation in the cultivated rice. Using functional markers to detect the genotype of GS3 and GW5 of 529 materials, through regression analysis, we found that two genes could explain more than 20% of the genetic variation of the stigma exposure, two The combination of genes can explain more than 30%. These results suggest that GS3 and GW5 are also the main gene.6. for controlling the exposure rate of stigma in cultivated rice. In order to study the feasibility of GS3 and GW5 for improved stigma exposure, we divide the two genes into four genotypes: GW5GS3, GW5gs3, gw5GS3 and gw5gs3. by comparing four genotypes. We found that the gene combination of GW5gs3 has the highest exposure rate and yield of stigma, and the highest exposure rate and yield of the genotype combination gw5GS3 in the cultivated rice are the lowest. Therefore, the genetic improvement of the cultivated rice with the dominant genotype combination of GW5gs3 is mainly in the presence of.7.GW5gs3 in the cultivated rice. In indica rice II and tropical japonica rice, gw5GS3 mainly exists in Indica I and temperate japonica rice. The difference of this distribution indicates that this GS3 and GW5 may be selected. Using these two gene sequences, SNPs analysis of the linkage disequilibrium of the two genes, we find that there is a strong linkage between the two genes whether or not the group structure is considered. Disequilibrium. Through the analysis of nucleic acid polymorphism of two genotypes and different genotypes, we found that these two genes may be selected in the process of breeding, thus forming differences in the distribution of GW5gs3 and gw5GS3 in different subgroups, leading to the difference in the exposure and yield of the stigma in different subgroups.
【學(xué)位授予單位】:華中農(nóng)業(yè)大學(xué)
【學(xué)位級別】:博士
【學(xué)位授予年份】:2017
【分類號】:S511

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