S20135420-何濤-染色體導(dǎo)入系在QTL定位中的應(yīng)用

染色體導(dǎo)入系在QTL定位中的應(yīng)用S20135420何濤摘要：染色體導(dǎo)入系是指借助分子輔助選擇方法，通過(guò)連續(xù)回交和自交構(gòu)建的只含有一個(gè)或幾個(gè)導(dǎo)入片段與受體親本不同，其他遺傳背景與受體親本完全相同的家系或品系。相對(duì)于傳統(tǒng)的遺傳群體，導(dǎo)入系群體大大降低了供體材料的遺傳背景的干擾，提高了QTL分析的靈敏度和準(zhǔn)確性，是QTL鑒定、精細(xì)定位、互作分析、圖位克隆、性狀改良及雜種優(yōu)勢(shì)利用和基因表達(dá)分析的理想的實(shí)驗(yàn)材料。本文對(duì)染色體導(dǎo)入系在QTL定位中的應(yīng)用進(jìn)行綜述。關(guān)鍵詞：染色體、導(dǎo)入系、QTL作物絕大多數(shù)經(jīng)濟(jì)性狀都屬于數(shù)量性狀，因此研究數(shù)量性狀的遺傳機(jī)制對(duì)于作物遺傳育種有非常重要的意義。而準(zhǔn)確鑒定和定位QTL是進(jìn)行數(shù)量性狀遺傳效應(yīng)分析、作物雜種優(yōu)勢(shì)利用研究以及分子克隆的基礎(chǔ)。但由于用于QTL鑒定和定位的傳統(tǒng)分析群體遺傳背景復(fù)雜，很難對(duì)單個(gè)QTL進(jìn)行準(zhǔn)確鑒定和定位[1]。所以一個(gè)良好的定位及分析群體是進(jìn)行數(shù)量性狀遺傳研究的必須條件，本文對(duì)染色體導(dǎo)入系在QTL定位中的研究情況進(jìn)行綜述。1QTLQTL（quantitativetraitlocus），即數(shù)量性狀基因座，指控制數(shù)量性狀的基因在基因組中的位置。QTL定位就是采用類(lèi)似單基因定位的方法將QTL定位在遺傳圖譜上，確定QTL與遺傳標(biāo)記間的距離（以重組率表示）。作物許多重要的農(nóng)藝性狀如品質(zhì)、產(chǎn)量和抗逆性等都屬于復(fù)雜數(shù)量性狀。作物QTL的不斷深入研究揭示了這些復(fù)雜數(shù)量性狀的遺傳基礎(chǔ)。促進(jìn)了這些性狀的遺傳改良，并使其分子剖析成為可能[2-4]。1.1QTL的定位群體QTL定位的精度在很大程度上取決于遺傳圖譜的標(biāo)記飽和度和分離群體的重組信息量。根據(jù)分離群體的特點(diǎn)，作圖群體分為:1.臨時(shí)性群體。如F2及其衍生的F3、F4家系，回交群體（BC）等。F2群體是常用的作圖群體，該群體容易配制且包含親本任意一個(gè)位點(diǎn)的所有基因型，更適合于估算基因效應(yīng)和檢測(cè)不同類(lèi)型的互作。但F2群體的一個(gè)不足之處是存在雜合基因型。對(duì)于顯性標(biāo)記，將無(wú)法識(shí)別顯性純合基因型和雜合基因型。F2群體的另一個(gè)缺點(diǎn)是不易長(zhǎng)期保存?；亟蝗后w每一分離的基因座只有兩種基因型，直接反映F1代配子的分離比例，因而其作圖效率最高，但缺點(diǎn)是不易保存。2.永久性分離群體。主要包括重組近交系（RILs）、雙單倍體群體（DH）、回交近交系（BILs）以及近等基因系（NILs）。它們共同的特點(diǎn)是系內(nèi)基因型是一致的，而系間基因型是不同的，系間除少數(shù)甚至一個(gè)染色體區(qū)段存在差異外，其余絕大部分染色體區(qū)段完全相同。因此，它們可以把種子分散到各個(gè)不同環(huán)境做重復(fù)試驗(yàn)以增加QTL檢測(cè)的準(zhǔn)確性，特別是NIL，由于消除了其他背景的干擾，甚至是主效QTL對(duì)微效QTL的掩蓋，因此QTL定位效率很高[5]。但也正由于這類(lèi)群體系內(nèi)基因型的一致性，因而不能估算顯性效應(yīng);而且，除非群體足夠大，否則它們提供的信息不如F2等群體。如DH群體，由于在染色體加倍過(guò)程中可能存在基因丟失，因此構(gòu)建分子標(biāo)記連鎖圖時(shí)盡量不用這種群體。1.2QTL定位分子標(biāo)記目前，研究所采用的分子標(biāo)記主要有基于分子雜交的RFLP（RestrictionFragmentLengthPolymorphism）[6]，基于PCR擴(kuò)增的RAPD（RandomAmplifiedPolymorphicDNA）[7]、SSR（SimpleSequenceRepeat）[8]、AFLP（AmplifiedFragmentLengthPolymorphism）[9]以及單核苷酸多態(tài)性標(biāo)記SNP（SingleNucleotidePolymorphism）[10]和表達(dá)序列標(biāo)簽EST（ExpressedSequenceTags）[11]等等。此外近年來(lái)還發(fā)展了序列標(biāo)志位點(diǎn)（SequencetaggedSites，STS）[12]、轉(zhuǎn)錄轉(zhuǎn)座子分子標(biāo)記（SequencespecificAmplificationPolymorphism，S-SAP）[13]、IRAP（InterretrotransposonAmplifiedPolymorphism）[14]、REMAP（RetrotransposonmicrosatelliteAmplificationPolymorphis）[15]和RBIP（RetrotransPoson-basedInsertionPolymorphism）[15]等。2染色體導(dǎo)入系染色體導(dǎo)入系(chromosomeintrogressionlines，CILs)，又稱(chēng)為染色體片段代換系(chromosomesegmentsubstitutionlines，CSSLs)或代換系(substitutionlines，SLs)，指只含有一個(gè)或幾個(gè)導(dǎo)入片段與受體親本不同，其他遺傳背景與受體親本完全相同的家系或品系。其中只含一個(gè)染色體片段與受體親本不同的，稱(chēng)為單片段代換系(singlesegmentsubstitutionline，SSSL)，這是最理想的代換系。相對(duì)于傳統(tǒng)的遺傳群體，導(dǎo)入系群體大大降低了供體材料的遺傳背景的干擾，提高了QTL分析的靈敏度和準(zhǔn)確性，是QTL鑒定、精細(xì)定位、互作分析、圖位克隆、性狀改良及雜種優(yōu)勢(shì)利用和基因表達(dá)分析的理想的實(shí)驗(yàn)材料。2.1染色體導(dǎo)入系的構(gòu)建染色體導(dǎo)入系的構(gòu)建主要是通過(guò)多代回交來(lái)建立。先將供體親本與受體親本雜交獲得F1，再以受體親本作為輪回親本，經(jīng)過(guò)多代回交并結(jié)合MAS技術(shù)篩選構(gòu)建大規(guī)模染色體片段導(dǎo)入系或代換系[16]。龍萍[17]利用我國(guó)和國(guó)際水稻所的21個(gè)優(yōu)良品種做輪回親本、來(lái)源于24個(gè)國(guó)家和地區(qū)的188份品種資源做供體親本，通過(guò)雜交、連續(xù)回交結(jié)合性狀篩選，構(gòu)建了近6萬(wàn)份具有輪回親本遺傳背景的近等基因?qū)胂?。Li等對(duì)2萬(wàn)個(gè)水稻ILs的定向選擇，鑒定得到了許多來(lái)源不同的新基因和耐旱、耐漬、特異的農(nóng)藝性狀和生理性狀的導(dǎo)入系，如273個(gè)耐旱ILs，175個(gè)耐鹽ILs和203個(gè)抗飛虱的ILs。2.2染色體導(dǎo)入系在QTL中的應(yīng)用目前主要農(nóng)作物中，水稻、玉米、棉花、大豆等均已建立了導(dǎo)入系或代換系群體，并在QTL定位中得到了大量應(yīng)用。在水稻上，Li等ADDINEN.CITE<EndNote><Cite><Author>Li</Author><Year>2011</Year><RecNum>238</RecNum><record><rec-number>238</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">238</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Li,Min</author><author>Sun,Penglin</author><author>Zhou,Hongju</author><author>Chen,Sheng</author><author>Yu,Sibin</author></authors></contributors><titles><title>Identificationofquantitativetraitlociassociatedwithgerminationusingchromosomesegmentsubstitutionlinesofrice(OryzasativaL.)</title><secondary-title>TAGTheoreticalandAppliedGenetics</secondary-title></titles><periodical><full-title>TAGTheoreticalandAppliedGenetics</full-title></periodical><pages>411-420</pages><volume>123</volume><number>3</number><keywords><keyword>BiomedicalandLifeSciences</keyword></keywords><dates><year>2011</year></dates><publisher>SpringerBerlin/Heidelberg</publisher><isbn>0040-5752</isbn><urls><related-urls><url>/10.1007/s00122-011-1593-9</url></related-urls></urls><electronic-resource-num>10.1007/s00122-011-1593-9</electronic-resource-num></record></Cite></EndNote>[18]在定位水稻發(fā)芽率的研究中，將粳稻品種Nipponbare的染色體片段導(dǎo)入珍秈97，得到了143個(gè)染色體代換系群體，在2、5、6、10號(hào)染色體上定位出4個(gè)相關(guān)QTL。再通過(guò)利用CSSL的F2群體，在2號(hào)染色體上證實(shí)了一個(gè)主效QTL(qGR2)，并從中找到一個(gè)在種子中優(yōu)先表達(dá)的候選基因OsMADS29。楊雷等ADDINEN.CITE<EndNote><Cite><Author>楊雷</Author><Year>2011</Year><RecNum>20</RecNum><record><rec-number>20</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">20</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>楊雷,</author><author>柳絮,</author><author>吳振映,</author><author>王文英,</author><author>姚方印,</author><author>劉開(kāi)啟,</author></authors></contributors><titles><title>水稻單片段代換系抽穗期qtl鑒定及遺傳分析</title><secondary-title>山東農(nóng)業(yè)科學(xué)</secondary-title><short-title>agri</short-title></titles><periodical><full-title>山東農(nóng)業(yè)科學(xué)</full-title></periodical><number>05</number><keywords><keyword>水稻</keyword><keyword>抽穗期</keyword><keyword>qtl鑒定</keyword><keyword>單片段代換系</keyword><keyword>遺傳分析</keyword></keywords><dates><year>2011</year></dates><isbn>1001-4942</isbn><work-type>Journal</work-type><urls></urls></record></Cite></EndNote>[19]以帶有晚抽穗基因的單片段代換系為供體，以華粳秈74為受體構(gòu)建單片段代換系，發(fā)展F2次級(jí)分離群體，定位出水稻第6染色體上存在與抽穗期相關(guān)的QTL，并鑒定出抽穗期由單基因控制，早抽穗表現(xiàn)為顯性。在此基礎(chǔ)上，WangADDINEN.CITE<EndNote><Cite><Author>Wang</Author><Year>2011</Year><RecNum>230</RecNum><record><rec-number>230</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">230</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Wang,Binbin</author><author>Zhu,Changxiang</author><author>Liu,Xu</author><author>Wang,Wenying</author><author>Ding,Hanfeng</author><author>Jiang,Mingsong</author><author>Li,Guangxian</author><author>Liu,Wei</author><author>Yao,Fangyin</author></authors></contributors><titles><title>FineMappingofqHD4-1,aQTLControllingtheHeadingDate,toa20.7-kbDNAFragmentinRice(OryzasativaL.)</title><secondary-title>PlantMolecularBiologyReporter</secondary-title></titles><periodical><full-title>PlantMolecularBiologyReporter</full-title></periodical><pages>702-713</pages><volume>29</volume><number>3</number><keywords><keyword>BiomedicalandLifeSciences</keyword></keywords><dates><year>2011</year></dates><publisher>SpringerNetherlands</publisher><isbn>0735-9640</isbn><urls><related-urls><url>/10.1007/s11105-010-0278-x</url></related-urls></urls><electronic-resource-num>10.1007/s11105-010-0278-x</electronic-resource-num></record></Cite></EndNote>[20]等、PeiADDINEN.CITE<EndNote><Cite><Author>Pei</Author><Year>2011</Year><RecNum>232</RecNum><record><rec-number>232</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">232</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Pei,Chengguo</author><author>Liu,Xu</author><author>Wang,Wenying</author><author>Ding,Hanfeng</author><author>Jiang,Mingsong</author><author>Li,Guangxian</author><author>Zhu,Changxiang</author><author>Wen,Fujiang</author><author>Yao,Fangyin</author></authors></contributors><titles><title>FineMappingofqHD8-1,aQTLControllingtheHeadingDate,toa26-kbDNAFragmentinRice(OryzasativaL.)</title><secondary-title>JournalofPlantBiology</secondary-title></titles><periodical><full-title>JournalofPlantBiology</full-title></periodical><pages>190-198</pages><volume>54</volume><number>3</number><keywords><keyword>BiomedicalandLifeSciences</keyword></keywords><dates><year>2011</year></dates><publisher>SpringerNewYork</publisher><isbn>1226-9239</isbn><urls><related-urls><url>/10.1007/s12374-011-9155-x</url></related-urls></urls><electronic-resource-num>10.1007/s12374-011-9155-x</electronic-resource-num></record></Cite></EndNote>[21]等，進(jìn)一步精細(xì)定位了qHD4-1和qHD8-1兩個(gè)QTL。趙芳明等ADDINEN.CITE<EndNote><Cite><Author>趙芳明</Author><Year>2011</Year><RecNum>21</RecNum><record><rec-number>21</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">21</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>趙芳明,</author><author>張桂權(quán),</author><author>曾瑞珍,</author><author>楊正林,</author><author>凌英華,</author><author>桑賢春,</author><author>何光華,</author></authors></contributors><titles><title>基于單片段代換系的水稻粒型qtl加性及上位性效應(yīng)分析</title><secondary-title>作物學(xué)報(bào)</secondary-title><short-title>xbzw</short-title></titles><periodical><full-title>作物學(xué)報(bào)</full-title></periodical><number>03</number><keywords><keyword>水稻</keyword><keyword>單片段代換系</keyword><keyword>粒型qtl</keyword><keyword>加性效應(yīng)</keyword><keyword>上位性效應(yīng)</keyword></keywords><dates><year>2011</year></dates><isbn>0496-3490</isbn><work-type>Journal</work-type><urls></urls></record></Cite></EndNote>[22]以l6個(gè)單片段代換系(SSSL)及15個(gè)雙片段代換系分析了水稻粒型性狀QTL的加性及上位性效應(yīng)，共檢測(cè)到9個(gè)水稻粒型性狀QTL和7對(duì)雙基因互作，揭示了同一粒長(zhǎng)QTL與不同單片段代換系聚合時(shí)會(huì)產(chǎn)生不同的互作效應(yīng)。李生強(qiáng)等ADDINEN.CITE<EndNote><Cite><Author>李生強(qiáng)</Author><Year>2011</Year><RecNum>17</RecNum><record><rec-number>17</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">17</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>李生強(qiáng),</author><author>崔國(guó)昆,</author><author>關(guān)成冉,</author><author>王俊,</author><author>梁國(guó)華,</author></authors></contributors><titles><title>基于水稻單片段代換系的粒形qtl定位</title><secondary-title>中國(guó)水稻科學(xué)</secondary-title><short-title>zgsk</short-title></titles><periodical><full-title>中國(guó)水稻科學(xué)</full-title></periodical><number>02</number><keywords><keyword>水稻</keyword><keyword>單片段代換系</keyword><keyword>粒形</keyword><keyword>數(shù)量性狀基因座</keyword></keywords><dates><year>2011</year></dates><isbn>1001-7216</isbn><work-type>Journal</work-type><urls></urls></record></Cite></EndNote>[23]利用22個(gè)單片段代換系，共鑒定出22個(gè)與粒形相關(guān)的QTL，包括7個(gè)粒長(zhǎng)QTL，6個(gè)粒寬QTL，5個(gè)谷粒長(zhǎng)寬比QTL和4個(gè)粒厚QTL。朱金燕等ADDINEN.CITE<EndNote><Cite><Author>朱金燕</Author><Year>2011</Year><RecNum>24</RecNum><record><rec-number>24</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">24</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>朱金燕,</author><author>嵇朝球,</author><author>周勇,</author><author>王軍,</author><author>王中德,</author><author>楊杰,</author><author>范方軍,</author><author>梁國(guó)華,</author><author>仲維功,</author></authors></contributors><titles><title>利用染色體單片段代換系定位水稻株高qtl</title><secondary-title>植物學(xué)報(bào)</secondary-title><short-title>zwxt</short-title></titles><periodical><full-title>植物學(xué)報(bào)</full-title></periodical><number>06</number><keywords><keyword>株高</keyword><keyword>數(shù)量性狀位點(diǎn)</keyword><keyword>水稻</keyword><keyword>單片段代換系</keyword><keyword>代換作圖</keyword></keywords><dates><year>2011</year></dates><isbn>1674-3466</isbn><work-type>Journal</work-type><urls></urls></record></Cite></EndNote>[24]以85個(gè)單片段代換系為材料定位了24個(gè)水稻株高QTL。任德勇等ADDINEN.CITE<EndNote><Cite><Author>任德勇</Author><Year>2010</Year><RecNum>18</RecNum><record><rec-number>18</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">18</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>任德勇,</author><author>何光華,</author><author>凌英華,</author><author>桑賢春,</author><author>楊正林,</author><author>趙芳明,</author></authors></contributors><titles><title>基于單片段代換系的水稻穗長(zhǎng)qtl加性及其上位性效應(yīng)</title><secondary-title>植物學(xué)報(bào)</secondary-title><short-title>zwxt</short-title></titles><periodical><full-title>植物學(xué)報(bào)</full-title></periodical><number>06</number><keywords><keyword>上位性效應(yīng)</keyword><keyword>穗長(zhǎng)</keyword><keyword>qtl</keyword><keyword>水稻</keyword><keyword>單片段代換系</keyword></keywords><dates><year>2010</year></dates><isbn>1674-3466</isbn><work-type>Journal</work-type><urls></urls></record></Cite></EndNote>[25]則利用染色體片段代換系進(jìn)行了水稻穗長(zhǎng)QTL加性及其上位性效應(yīng)分析。在棉花上，WangP等ADDINEN.CITEADDINEN.CITE.DATA[26]以海島棉（G.barbadense）TM-1為受體親本，陸地棉(G.barbadense)Hai7124為供體親本，構(gòu)建染色體片段導(dǎo)入系（CHSLs）。該群體由包含298個(gè)導(dǎo)入片段的174個(gè)群體組成，其中86個(gè)群體（49.1%）為單片段導(dǎo)入。導(dǎo)入片段平均長(zhǎng)度為16.7cM，總長(zhǎng)度為2948.7cM，覆蓋了83.3%的4倍體棉花基因組。利用此群體，通過(guò)對(duì)4個(gè)環(huán)境下收集的數(shù)據(jù)進(jìn)行綜合分析，發(fā)現(xiàn)了與棉花纖維質(zhì)量相關(guān)的43個(gè)加性效應(yīng)QTL和6個(gè)上位性效應(yīng)QTL，其中有6個(gè)QTL在各環(huán)境中均表現(xiàn)穩(wěn)定。在玉米上，Chung等ADDINEN.CITE<EndNote><Cite><Author>Chung</Author><Year>2010</Year><RecNum>255</RecNum><record><rec-number>255</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">255</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Chung,C.L.</author><author>Longfellow,J.M.</author><author>Walsh,E.K.</author><author>Kerdieh,Z.</author><author>VanEsbroeck,G.</author><author>Balint-Kurti,P.</author><author>Nelson,R.J.</author></authors></contributors><auth-address>Dept.ofPlantPathologyandPlant-MicrobeBiology,CornellUniversity,Ithaca,NY14853,USA.</auth-address><titles><title>Resistancelociaffectingdistinctstagesoffungalpathogenesis:useofintrogressionlinesforQTLmappingandcharacterizationinthemaize--Setosphaeriaturcicapathosystem</title><secondary-title>BMCPlantBiol</secondary-title></titles><periodical><full-title>BMCPlantBiol</full-title></periodical><pages>103</pages><volume>10</volume><edition>2010/06/10</edition><keywords><keyword>Ascomycota/physiology</keyword><keyword>ChromosomeMapping</keyword><keyword>DNA,Plant/genetics</keyword><keyword>Host-PathogenInteractions</keyword><keyword>Immunity,Innate</keyword><keyword>Models,Genetic</keyword><keyword>Phenotype</keyword><keyword>PlantDiseases/genetics</keyword><keyword>QuantitativeTraitLoci</keyword><keyword>Zeamays/genetics/immunology/microbiology</keyword></keywords><dates><year>2010</year></dates><isbn>1471-2229(Electronic) 1471-2229(Linking)</isbn><accession-num>20529319</accession-num><urls></urls><custom2>3017769</custom2><electronic-resource-num>1471-2229-10-103[pii] 10.1186/1471-2229-10-103[doi]</electronic-resource-num><remote-database-provider>Nlm</remote-database-provider><language>eng</language></record></Cite></EndNote>[27]用Tx303和B73為親本構(gòu)建的82個(gè)代換系群體對(duì)玉米大斑病進(jìn)行QTL定位和分析。得到了兩個(gè)可靠表達(dá)的QTL：qNLB1.06和qNLB1.02。在溫室與田間的重復(fù)試驗(yàn)表明：qNLB1.06減少真菌侵入的效率，qNLB1.02增加侵染位點(diǎn)附近胼胝質(zhì)和多酚的積累，阻礙菌絲向維管束中生長(zhǎng)。除了對(duì)玉米大斑病的抗性，qNLB1.02還與玉米細(xì)菌性枯萎病和普通銹病有關(guān)，而qNLB1.06也增強(qiáng)對(duì)玉米細(xì)菌性枯萎病的抗性。EvertonA.Brenner等ADDINEN.CITE<EndNote><Cite><Author>Brenner</Author><Year>2012</Year><RecNum>236</RecNum><record><rec-number>236</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">236</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Brenner,Everton</author><author>Blanco,Mike</author><author>Gardner,Candice</author><author>Lübberstedt,Thomas</author></authors></contributors><titles><title>Genotypicandphenotypiccharacterizationofisogenicdoubledhaploidexoticintrogressionlinesinmaize</title><secondary-title>MolecularBreeding</secondary-title></titles><periodical><full-title>MolecularBreeding</full-title></periodical><pages>1-16</pages><keywords><keyword>BiomedicalandLifeSciences</keyword></keywords><dates><year>2012</year></dates><publisher>SpringerNetherlands</publisher><isbn>1380-3743</isbn><urls><related-urls><url>/10.1007/s11032-011-9684-5</url></related-urls></urls><electronic-resource-num>10.1007/s11032-011-9684-5</electronic-resource-num></record></Cite></EndNote>[28]將31個(gè)外源玉米種質(zhì)導(dǎo)入PHZ51和PB47中，得到50個(gè)BC1DH群體，含有199個(gè)SNP位點(diǎn)分布在整個(gè)基因組，其中平均11.8%的標(biāo)記來(lái)自于外源供體等位基因。這50個(gè)BC1DH群體包含了輪回親本92.9%的基因組。利用此群體，通過(guò)關(guān)聯(lián)分析揭示了細(xì)胞壁消化、倒伏和花期推遲有關(guān)的數(shù)量性狀多態(tài)性。邢向茹等ADDINEN.CITE<EndNote><Cite><Author>邢向茹</Author><Year>2011</Year><RecNum>37</RecNum><record><rec-number>37</rec-number><foreign-keys><keyapp="EN"db-id="9pwdta05da9waiedtz2vxtde0t2w0r0r50fa">37</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>邢向茹,</author><author>陳晨,</author><author>劉瑞響,</author><author>趙璞,</author><author>李成璞,</author><author>張祖新,</author></authors></contributors><titles><title>低氮脅迫對(duì)玉米染色體片段導(dǎo)入系產(chǎn)量和苗期性狀的影響</title><secondary-title>安徽農(nóng)業(yè)科學(xué)</secondary-title><short-title>ahny</short-title></titles><periodical><full-title>安徽農(nóng)業(yè)科學(xué)</full-title></periodical><number>21</number><keywords><keyword>玉米</keyword><keyword>氮脅迫</keyword><keyword>產(chǎn)量</keyword><keyword>苗期性狀</keyword><keyword>處理時(shí)期</keyword></keywords><dates><year>2011</year></dates><isbn>0517-6611</isbn><work-type>Journal</work-type><urls></urls></record></Cite></EndNote>[29]以Ye478及Ye478為背景的5個(gè)染色體片段導(dǎo)人系為材料，研究了低氮脅迫下產(chǎn)量和苗期性狀的影響。展望理想狀態(tài)下，整個(gè)導(dǎo)入系覆蓋了受體親本的全基因組，不同導(dǎo)入系帶有供體親本基因組不同片段。與輪回親本比較，兩個(gè)株系之間性狀的任何顯著差異都是因?yàn)閷?dǎo)入片段在供體與受體間存在的等位差異，因而可有效消除基因組上其他位點(diǎn)因分離而產(chǎn)生的遺傳差異，提高QTL定位的準(zhǔn)確性和穩(wěn)定性。染色體導(dǎo)入系為更深入研究QTL提供了良好的條件，特別是染色體單片段代換系，在QTL的鑒定和精細(xì)定位，QTL遺傳效應(yīng)分析及QTL克隆以及應(yīng)用于品種改良和雜種優(yōu)勢(shì)利用的研究中具有巨大的優(yōu)越性，為我們的研究帶來(lái)極大的方便。參考文獻(xiàn)[1].DarvasiA.,WeinrebA.,MinkeV.,WellerJ.I.,andSollerM..Detectingmarker-QTLlinkageandestimatingQTLgeneeffectandmaplocationusingasaturatedgeneticmap.Genetics,1993,134:943-951.[2].ZhuH,HayesP,KleinhofsA,KudrnaD,LiuZ,PromL,SteffensonB,ToojindaT,VivarH,GilchristL,1999.DoesfunctionfollowformPrincipalQTLsforFusariumheadblight(FHB)resistancearecoincidentwithQTLsforinflorescencetraitsandplantheightinadoubled-haploidpopulationofbarley.TheoreticalandAppliedGenetics,99(7～8):1221～1232[3].YanJQ,HeCX,BenmoussaM,WuP,ZhuJ,1998.Quantitativetraitlocianalysisforthedevelopmentalbehavioroftillernumberinrice(OryzasativaL.).TheoreticalandAppliedGenetic,97(1～2):267～274[4].YanJQ,ZhouJ,HeCX,MebroukB,WuP,1998b.Moleculardissectionofdevelopmentalbehaviorofplantheightinrice(OryzasativaL.).Genetics,150:1257～1265[5].朱軍.運(yùn)用混合線性模型定位復(fù)雜數(shù)量性狀基因的方法J.浙江大學(xué)學(xué)報(bào),1999,333:327–3351[6].GRODZICKERT,WIILIAMSJ.SHARPP.SAMBROOKJ.Physicalmappingoftemperature-sensitivemutationsofadenoviruses[J].ColdSpringHarbor.Syrup.Quant.Biol.1974.39：439—446.[7].WILLIAMSJGK,KUBEIIKAR,LIVAKKJ,etal.DNApolymorphismsamplifiedbyarbitraryprimersareusefulasgeneticmarkers[J].Nucleic.Acids.Res.,1990,18：6531-6535.[8].AKKAYAMS.BHAGWATAA.CREGANPB.LengthpolymorphismsofsimplesequencerepeatDNAinsoybean[J].Genera,1992,132：1131-1139.[9].VOSP,HOGERSR,BLEEKERM,eta1.AFIP：AnewtechniqueforDNAfingerprinting[J].Nucl.Acid.Res.1995,23：4407-4414.[10].CHORJ,MINDRINOSM,RICHARDSDR.ela1.GenomewidemappingwithbiallelicmarkersinArabidopsisthaliana[J].NatureGenetics。1999,23：203-207.[11].ADAMSMD,KEIIEYJM,GOCAYNEJD,elal.ComplementaryDNAsequencing：expressedsequencetagsandhumangenomeproject[J].Science,1991,252(5013)：1651-1656.[12].OLSONM,HOODL,CANTORC,etal.Acommonlanguageforphysicalmappingofthehumangenome[J].SciemP,1989,245(4925)：1434-1435.[13].WAUGHR,MCLEANK,FIAVEI1AJ,elal.GeneticdistributionofBARE-1-likeretrotransposableelementsint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