大豆α-生育酚的遗传与QTL分析
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摘要
【目的】通过对大豆α-生育酚进行遗传和QTL分析,研究其遗传机制,定位其主效QTL,为高α-生育酚含量的大豆品种选育奠定遗传学基础。【方法】以栽培大豆晋豆23为母本、山西农家品种大豆灰布支黑豆(ZDD02315)为父本杂交衍生的447个RIL作为供试群体构建遗传图谱,试验群体及亲本分别于2011年、2012年和2015年夏季在河南省农业科学院原阳试验基地种植,冬季在海南省三亚南繁基地种植。田间试验采取随机区组设计,2次重复。从6个环境中每个家系选取15.00 g籽粒饱满,大小一致的大豆种子,利用高效液相色谱法定性、定量测定样品中的α-生育酚含量。采用主基因+多基因混合遗传分离分析法和WinQTLCart 2.5复合区间作图法,对大豆α-生育酚含量进行主基因+多基因混合遗传分析和QTL定位。【结果】基于主基因+多基因混合遗传分离分析法,α-生育酚受4对主基因控制,遗传基因分布在双亲中。4对主基因间加性效应值中3对为正值,表明这些基因来源于母本晋豆23;1对为负值,表明该对基因来源于父本灰布支黑豆;4对主基因之间相互作用的上位性效应表现为正值和负值的各有3对,说明不同基因间上位性效应对α-TOC的影响方向并不完全一致。环境因素引起的变异为0.13%—4.05%。表明α-TOC主要受4对主基因影响,受环境因素影响较小。采用WinQTLCart 2.5复合区间作图(CIM)共检测到17个影响α-生育酚的QTL,分布于第1、2、5、6、8、14、16、17共8条染色体中,单个QTL的贡献率8.35%—35.78%,QTL主要表现为加性效应。qα-D1a-1同时在2011年原阳、2012年原阳和三亚、2015年原阳4个环境下检测到,且均定位在第1染色体Satt320—Satt254标记区间19.79 cM处,解释的表型变异分别为12.55%、12.01%和11.89%、12.61%,加性效应值0.119-0.132,增加α-TOC含量的等位基因来自母本晋豆23;qα-A2-1同时在2011年原阳和三亚、2015年原阳3个环境下检测到,且均定位在第8染色体Sat_129—Satt377标记区间44.53 cM处,解释的表型变异分别为23.18%和22.56%、23.01%,加性效应值-0.195—-0.180,增加α-TOC含量的等位基因来自父本灰布支黑豆。qα-D1a-1和qα-A2-1 2个QTL能够稳定遗传。【结论】α-生育酚最适遗传模型符合4MG-AI,即4对具有加性上位性效应的主基因遗传模型。其遗传主要受4对主基因影响,受环境因素影响较小。检测到α-生育酚的2个稳定主效QTL,Satt320—Satt254和Sat_129—Satt377是共位标记区间。
【Objective】 Inheritance and main QTL for α-tocopherol were detected by genetic analysis and QTL mapping. The results lay a genetic foundation for the selection of soybean varieties with high α-tocopherol content in soybean.【Method】The 447 RILs were derived from a cross between Jindou23 of commercial cultivar as the female parent and Huibuzhi of farm variety from Shanxi Province(ZDD02315) as the male parent that construct SSR genetic linkage map. The parent lines and the RILs were cultivated in summer at Yuanyang testing ground of Academy of Agricultural Sciences, and in Winter at Sanya of Hainan province in 2011, 2012, 2015. A complete random design with two replications was used in this study. Each plot of a single genotype provided 15.00 g big-plump seeds with same size in six environmental conditions. α-tocopherol content was detected quantitatively and qualitatively by High Performance Liquid Chromatography(HPLC). Major gene plus polygene mixed inheritance and QTL mapping for α-tocopherol were detected by major gene plus polygene mixed inheritance analysis and composite interval mapping with WinQTLCart 2.5.【Result】The results showed that α-tocopherol was controlled by four pairs of main genes by major gene plus polygene mixed inheritance analysis. and the four pairs of main genes distributed in two parents. Three main genes shared the same direction with positive additive effect and involved novel alleles from the same parent, Jindou23; one main gene has negative additive effects and donated by Huibuzhi of black beans. Three pairs of genes shared the different direction of positive or negative epistatic effects shared the different direction to α-tocopherol contribution. The phenotypic variation explained by QTL by environment interaction ranged from 0.13 to 4.05%, and indicated that α-tocopherol was significantly affected by four pairs of main genes, more than by environment. Seventeen QTLs for α-tocopherol were mapped on 8 chromosomes 1, 2, 5, 6, 8, 14, 16, and 17, separately; the variation accounted for by each of these seventeen QTLs ranged from 8.35% to 35.78%; and QTL showed additive effect. qα-D1 a-1 was all located in marker intervals between Satt320-Satt254(19.79 cM) on chromosomes 1 in four environmental conditions of 2011 at Yuanyang, 2012 at Yuanyang and Sanya, and 2015 at Yuanyang. and explained 12.55%, 12.01%, 11.89%, 12.61% of the phenotypic variation. It had an additive effect of 0.119-0.132 donated by Jinbean23. qα-A2-1 was all located in marker intervals between Sat_129-Satt377(44.53 cM) on chromosomes 8 in three environmental conditions of 2011 at Yuanyang and Sanya, 2015 at Yuanyang. and explained 23.18%, 22.56%, 23.01% of the phenotypic variation. It had an additive effect of-0.195--0.180 donated by Huibuzhi. qα-D1 a-1 and qα-A2-1 can be stably expressed in different genetic backgrounds. 【Conclusion】α-tocopherol was controlled by four pairs of additive epistatic effect major genes genetic model(4 MG-AI), and it less affected by environmental factor. The two stable main QTL of Satt320-Satt254 and Sat_129-Satt377 were co-localization marker intervals in soybean.
【Objective】 Inheritance and main QTL for α-tocopherol were detected by genetic analysis and QTL mapping. The results lay a genetic foundation for the selection of soybean varieties with high α-tocopherol content in soybean.【Method】The 447 RILs were derived from a cross between Jindou23 of commercial cultivar as the female parent and Huibuzhi of farm variety from Shanxi Province(ZDD02315) as the male parent that construct SSR genetic linkage map. The parent lines and the RILs were cultivated in summer at Yuanyang testing ground of Academy of Agricultural Sciences, and in Winter at Sanya of Hainan province in 2011, 2012, 2015. A complete random design with two replications was used in this study. Each plot of a single genotype provided 15.00 g big-plump seeds with same size in six environmental conditions. α-tocopherol content was detected quantitatively and qualitatively by High Performance Liquid Chromatography(HPLC). Major gene plus polygene mixed inheritance and QTL mapping for α-tocopherol were detected by major gene plus polygene mixed inheritance analysis and composite interval mapping with WinQTLCart 2.5.【Result】The results showed that α-tocopherol was controlled by four pairs of main genes by major gene plus polygene mixed inheritance analysis. and the four pairs of main genes distributed in two parents. Three main genes shared the same direction with positive additive effect and involved novel alleles from the same parent, Jindou23; one main gene has negative additive effects and donated by Huibuzhi of black beans. Three pairs of genes shared the different direction of positive or negative epistatic effects shared the different direction to α-tocopherol contribution. The phenotypic variation explained by QTL by environment interaction ranged from 0.13 to 4.05%, and indicated that α-tocopherol was significantly affected by four pairs of main genes, more than by environment. Seventeen QTLs for α-tocopherol were mapped on 8 chromosomes 1, 2, 5, 6, 8, 14, 16, and 17, separately; the variation accounted for by each of these seventeen QTLs ranged from 8.35% to 35.78%; and QTL showed additive effect. qα-D1 a-1 was all located in marker intervals between Satt320-Satt254(19.79 cM) on chromosomes 1 in four environmental conditions of 2011 at Yuanyang, 2012 at Yuanyang and Sanya, and 2015 at Yuanyang. and explained 12.55%, 12.01%, 11.89%, 12.61% of the phenotypic variation. It had an additive effect of 0.119-0.132 donated by Jinbean23. qα-A2-1 was all located in marker intervals between Sat_129-Satt377(44.53 cM) on chromosomes 8 in three environmental conditions of 2011 at Yuanyang and Sanya, 2015 at Yuanyang. and explained 23.18%, 22.56%, 23.01% of the phenotypic variation. It had an additive effect of-0.195--0.180 donated by Huibuzhi. qα-D1 a-1 and qα-A2-1 can be stably expressed in different genetic backgrounds. 【Conclusion】α-tocopherol was controlled by four pairs of additive epistatic effect major genes genetic model(4 MG-AI), and it less affected by environmental factor. The two stable main QTL of Satt320-Satt254 and Sat_129-Satt377 were co-localization marker intervals in soybean.
引文
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[4]KANWISCHER M,PORFIROVA S,BERGMULLER E,D?RMANN P.Alterations in tocopherol cyclase activity in transgenic and mutant plants of Arabidopsis affect tocopherol content,tocopherol composition,and oxidative stress.Plant Physiology,2005,137:713-723.
[5]TAVVA V S,KIM Y H,KAGAN I A,DINKINS R D,KIM K H,COLLINS G B.Increased alpha-tocopherol content in soybean seed overexpressing the Perilla frutescens gamma-tocopherol methyltransferase gene.Plant Cell Reports,2007,26(1):1-70.
[6]李海燕,隋美楠,聂腾坤,史帅,韩英鹏,李文滨.大豆维生素E五个相关基因表达模式分析.东北农业大学学报,2016,47(5):15-22.LI H Y,SUI M N,NIE T K,SHI S,HAN Y P,LI W B.Expression analysis of five relative genes of soybean vitamin E.Journal of Northeast Agricultural University,2016,47(5):15-22.(in Chinese)
[7]FENG F,DENG F,ZHOU P,YAN J,WANG Q,YANG R,LI X.QTLmapping for the tocopherols at milk stage of kernel development in sweet corn.Euphytica,2013,193(3):409-417.
[8]XU S,ZHANG D,CAI Y,ZHOU Y,TRUSHAR S,FARHAN A,LI Q,LI Z,WANG W,LI J,YANG X,YAN J.Dissecting tocopherols content in maize(Zea mays L.),using two segregating populations and high-density single nucleotide polymorphism markers.BMCPlant Biology,2012,12(1):201.
[9]WANG X,ZHANG C,LI L,FRITSCHE S,ENDRIGKEIT J,ZHANG W,LONG Y,JUNG C,MENG J.Unraveling the genetic basis of seed tocopherol content and composition in rapeseed(Brassica napus L.).PLoS ONE,2012,7(11):e50038.
[10]GRAEBNER R C,WISE M,CUESTA-MARCOS A,GENIZA M,BLAKE T,BLAKE V C,BUTLER J,CHAO S,HOLE D J,HORSLEY R,JAISWAL P,OBERT D,SMITH K,ULLRICHL S,HAYESL P M.Quantitative trait loci associated with the tocochromanol(vitamin E)pathway in barley.PLoS ONE,2015,10(7):e0133767.
[11]HADDADI P,EBRAHIMI A,LANGLADE N B,YAZDI-SAMADI B,BERGER M,CALMON A,NAGHAVI M R,VINCOURT P,SARRAFI A.Genetic dissection of tocopherol and phytosterol in recombinant inbred lines of sunflower through quantitative trait locus analysis and the candidate gene approach.Molecular Breeding,2012,29(3):717-729.
[12]MORAL L D,FERNANDEZ-MARTINEZ J M,VELASCO L,PEREZVICH.Quantitative trait loci for seed tocopherol content in sunflower.Crop Science,2012,52(2):786-794.
[13]GUPTA S,SANGHA M K,KAUR G,BANGA S,GUPTA M,KUMAR H,BANGA S S.QTL analysis for phytonutrient compounds and the antioxidant molecule in mustard(Brassica juncea L.).Euphytica,2015,201(3):345-356.
[14]DWIYANTI M S,UJIIE A,THUY L T B,YAMDA T,KITAMURA K.Genetic analysis of highα-tocopherol content in soybean seeds.Breed Science,2007,57:23-28.
[15]SHAW E,RAJCAN I.Molecular mapping of soybean seed tocopherols in the cross‘OAC Bayfield’בOAC Shire’.Plant Breeding,2017,136:83-93.
[16]LI H,WANG Y,HAN Y,TENG W,ZHAO X,LI Y,LI W.Mapping quantitative trait loci(QTLs)underlying seed vitamin E content in soybean with main,epistatic and QTL×environment effects.Plant Breeding,2016,135(2):208-214.
[17]LI H,LIU H,HAN Y,WU X,TENG W,LIU G,LI W.Identification of QTL underlying vitamin E contents in soybean seed among multiple environments.Theoretical and Applied Genetics,2010,120(7):1405-1413.
[18]张红梅,李海朝,文自翔,顾和平,袁星星,陈华涛,崔晓艳,陈新,卢为国.大豆籽粒维生素E含量的QTL分析.作物学报,2015,41(2):187-196.ZHANG H M,LI H C,WEN Z X,GU H P,YUAN X X,CHEN H T,CUI X Y,CHEN X,LU W G.Identification of QTL associated with vitamin E content in soybean seeds.Acta Agronomica Sinica,2015,41(2):187-196.
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