大豆γ-生育酚的混合遗传分析与QTL定位

doi:10.3864/j.issn.0578-1752.2020.11.002

摘要/Abstract

摘要：

【目的】通过对大豆γ-生育酚进行混合遗传和QTL定位分析,了解其遗传机制,定位其主效QTL,为高γ-生育酚含量大豆品种的选育奠定基础。【方法】以栽培大豆晋豆23为母本,以山西农家品种大豆灰布支黑豆为父本杂交衍生的重组自交系作为供试群体构建遗传图谱。图谱全长2 047.6 cM,平均图距8.8 cM,包括227个SSR标记,232个标记位点。重组自交系试验群体及亲本材料分别于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试验基地种植。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γ-生育酚含量。采用主基因+多基因混合遗传分离分析法,对大豆γ-生育酚含量进行混合遗传分析;采用WinQTLCart 2.5复合区间作图法,对大豆γ-生育酚含量进行QTL定位分析。【结果】主基因+多基因混合遗传分析表明,γ-生育酚受2对重叠作用主基因×加性多基因控制。遗传基因分布在双亲中。三亚试验数据检测出2对主基因间上位性效应值为0.4010—0.5169和多基因的加性效应值为0.1797—0.2146,主基因遗传率为11.27%—13.05%,多基因遗传率为82.51%—86.55%,多基因效应大于主基因效应。原阳试验数据检测到2对主基因间上位性效应值为0.9646—1.8455,主基因遗传率为39.51%—58.96%,没有检测出多基因效应。采用WinQTLCart 2.5复合区间作图（CIM）共检测到9个影响γ-生育酚的QTL,分布于A1（Chr.5）、A2（Chr.8）、C1（Chr.4）、K（Chr.9）、M（Chr.7）和G（Chr.8）6条染色体中,单个QTL的贡献率为7.29%—29.55%。qγ-G-1同时在2011年原阳、2012年三亚、2015年三亚3个环境下检测到,且均定位在G（Chr.18）染色体Satt275—Satt038标记区间0.01 cM处,解释的表型变异分别为8.97%、8.12%和7.91%。qγ-A1-1同时在2011年原阳和2015年原阳2个环境下检测到,且均定位在A1（Chr.5）染色体Satt276—Satt364标记区间0.01 cM处,解释的表型变异分别为29.54%和28.23%。qγ-G-1和qγ-A1-1 2个QTL能够稳定遗传。【结论】γ-T最适遗传模型符合MX2-Duplicate-A,即2对重叠作用主基因×加性多基因模型。其遗传同时受到基因型、环境和上位性的影响。检测到γ-T的2个稳定主效QTL,Satt275—Satt038和Satt276—Satt364是共位标记区间。

关键词: 大豆, γ-生育酚, 遗传, 主基因+多基因, 基因定位, 上位性

Abstract:

【Objective】Inheritance and main QTL for the content of γ-tocopherol were detected by genetic analysis and QTL mapping. The results lay a genetic foundation for the selection of soybean varieties with high γ-tocopherol content in soybean. 【Method】The RILs were derived from a cross between Jindou23 of commercial cultivar as the female parent and Huibuzhi of farm variety from Shanxi Province as the male parent that construct genetic linkage map. The map consisting of 232 marker loci spanned a total of 2 047.6 cM in length with an average spacing of 8.8 cM between adjacent marker loci. The parent lines and the RILs were cultivated in summer at Yuanyang experimental farm of Henan Academy of Agricultural Sciences, and in winter at Sanya of Hainan province in 2011, 2012, 2015. Random block design was adopted in field experiment, and the entire planting experiment was replicated twice. 15.00 g fully filled and uniform soybean seeds from each RILs in six environments were sampled. The content of γ-tocopherol was quantitatively and qualitatively analysis by High Performance Liquid Chromatography (HPLC). The content of γ-tocopherol in soybean was analyzed by major gene plus polygene mixed inheritance model approach. QTL for the content of γ-tocopherol in soybean were detected by composite interval mapping model using WinQTLCart 2.5. 【Result】The results showed that the content of γ-tocopherol was controlled by two pairs of main overlapping major gene × additive polygenes using major gene plus polygene mixed inheritance analysis. According to the data of Sanya, the epistasis effect value between two major genes was 0.4010-0.5169, and the additive effect value of polygene was 0.1797-0.2146. The results of Sanya experiment showed that the heritability of major gene and polygene were 11.27%-13.05% and 82.51%-86.55%, respectively. The polygene effect was greater than that of major gene effect. The data of Yuanyang experiment showed that the epistasis effect between the two major genes was 0.9646-1.8455, and the heritability of the major genes was 39.51%-58.96%. No polygenic effect was detected. QTLs resolved by using WinQTLCart 2.5 Compound Interval Mapping (CIM) analysis. Nine QTLs for the content of γ-tocopherol were mapped on chromosomes A1(Ch.5), A2(Chr.8), C1(Chr.4), K(Chr.9), M(Chr.7) and G(Chr.18), respectively. These QTL individually explained 7.29%-29.55% of the total phenotypic variation. The QTL of qγ-G-1 flanked by Satt275 and Satt038 (0.01 cM) on chromosome 8, were detected in three environmental conditions of 2011 at Yuanyang, 2012 and 2015 at Sanya, and explained 8.97%, 8.12%, 7.91% of the phenotypic variation, respectively. The QTL of qγ-A1-1 flanked by Satt276 and Satt364 (0.01 cM) on chromosomes 5, were detected in three environmental conditions of 2011 and 2015 at Yuanyang, and explained 29.54%, 28.23% of the phenotypic variation, respectively. qγ-G-1 and qγ-A1-1 can be stably expressed in different genetic backgrounds. 【Conclusion】The content of γ-tocopherol was controlled by two pairs of overlapping major Gene × additive Polygenes genetic model (MX2-Duplicate-A). Its inheritance was influenced by gene, environment, and epistasis. The two stable inheritance main-effect QTL for the content of γ-tocopherol were co-localization with marker Satt275-Satt038 and Satt276-Satt364 intervals in soybean, respectively. The co-localization marker interval has certain reference value for molecular marker assisted soybean breeding.

Key words: soybean, γ-Tocopherol, genetic, major genes plus polygene, gene mapping, epistasis

梁慧珍,许兰杰,董薇,余永亮,杨红旗,谭政委,李磊,刘新梅. 大豆γ-生育酚的混合遗传分析与QTL定位[J]. 中国农业科学, 2020, 53(11): 2149-2160.

LIANG HuiZhen,XU LanJie,DONG Wei,YU YongLiang,YANG HongQi,TAN ZhengWei,LI Lei,LIU XinMei. Mixed Inheritance Analysis and QTL Mapping for γ-Tocopherol Content in Soybean[J]. Scientia Agricultura Sinica, 2020, 53(11): 2149-2160.

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参考文献 49

[1]	RIMBACH G, MOEHRING J, HUEBBE P . Gene-regulatory activity of alpha-tocopherol. Molecules, 2010,15:1746-1761. doi: 10.3390/molecules15031746
[2]	ABBASI A R, HAJIREZAEI M, HOFIUS D, SONNEWALD U, VOLL L M . Specific roles of α- and γ-tocopherol in abiotic stress responses of transgenic tobacco. Plant Physiology, 2007,143:1720-1738. doi: 10.1104/pp.106.094771
[3]	HINCHA D K . Effects of α-tocopherol (Vitamin E) on the stability and lipid dynamics of model membranes mimicking the lipid composition of plant chloroplast membranes. FEBS Letters, 2008,582:3687-3692.
[4]	吴雪辉, 蓝梧涛, 容欧 . 响应面优化油茶籽油脱臭馏出物中维生素E的提取工艺研究. 中国油脂, 2018,43(5):113-116.
	WU X H, LAN W T, RONG O . Optimization of extraction of vitamin E from deodorizer distillate of oil-tea camellia seed oil by response surface methodology. China Oil and Fats, 2018,43(5):113-116. (in Chinese)
[5]	王哲, 毛善俊, 李浩然, 王勇 . 维生素E的催化合成路线分析. 物理化学学报, 2018,34(6):598-617.
	WANG Z, MAO S J, LI H R, WANG Y . How to synthesize vitamin E. Acta Physico-Chimica Sinica, 2018,34(6):598-617. (in Chinese)
[6]	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. doi: 10.1007/s00299-006-0207-5
[7]	KANWISCHER M, PORFIROVA S, BERGMULLER E . 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.
[8]	李辉, 罗非君, 林亲录 . γ-生育酚抗癌作用研究进展. 粮食与油脂, 2013,26(3):6-8.
	LI H, LUO F J, LIN Q L . Research progress on anticancer of gamma-tocopherol. Cerea1s & Oils, 2013,26(3):6-8. (in Chinese)
[9]	郑景洲, 许友卿, 丁兆坤 . 维生素E的功能、吸收与代谢(1). 生物学通报, 2005,40(11):1-2.
	ZHENG J Z, XU Y Q, DING Z K . Function, absorption and metabolism of vitamin E (1). Bulletin of Biology, 2005,40(11):1-2. (in Chinese)
[10]	SONTAG T J . Cytochrome P450 omega-hydroxylase pathway of tocopherol catabolism. novel mechanism of regulation of vitamin E status. Journal of Biological Chemistry, 2002,277:25290-25296. doi: 10.1074/jbc.M201466200
[11]	JIANG Q, RAO X, KIM C Y, FREISER H, ZHANG Q, JIANG Z Y, LI G L . Gamma-tocotrienol induces apoptosis and autophagy in prostate cancer cells by increasing intracellular dihydrosphingosine and dihydroceramide. International Journal of Cancer, 2012,130(3):685-693.
[12]	HUANG Y, KHOR T O, SHU L M, SAW C L, WU T Y, SUH N, YANG C S, KONG A N T . A γ-tocopherol-rich mixture of tocopherols maintains Nrf 2 expression in prostate tumors of TRAMP mice via epigenetic inhibition of CpG methylation. The Journal of Nutrition, 2012,142(5):818-823.
[13]	YANG C S, SUH N . Cancer prevention by different forms of tocopherols. Topics in Current Chemistry, 2013,329:21-33.
[14]	FENG F, DENG F, ZHOU P, YAN J, WANG Q, YANG R, LI X . QTL mapping for the tocopherols at milk stage of kernel development in sweet corn. Euphytica, 2013,193(3):409-417. doi: 10.1007/s10681-013-0948-5
[15]	GUPTA S, SANGHA M K, KAUR G, BANGA S S, BANGA S S . QTL analysis for phytonutrient compounds and the antioxidant molecule in mustard (Brassica juncea L.). Euphytica, 2015,201(3):345-356.
[16]	HADDADI P, EBRAHIMI A, LANGLADE N B, YAZDI-SAMADI B, BERGER M, CALMON 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. doi: 10.1007/s11032-011-9585-7
[17]	GRAEBNER R C, WISE M, CUESTA-MARCOS A, GENIZA M, BLAKE T, BLAKE V C . Quantitative trait loci associated with the tocochromanol (vitamin E) pathway in barley. PLoS ONE, 2015,10(7):e0133767.
[18]	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.
[19]	LI H, WANG Y, HAN Y, TENG W, ZHAO X, LI Y . 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. doi: 10.1111/pbr.2016.135.issue-2
[20]	LI H, LIU H, HAN Y, WU X, TENG W, LIU G . Identification of QTL underlying vitamin E contents in soybean seed among multiple environments. Theoretical and Applied Genetics, 2010,120(7):1405-1413. doi: 10.1007/s00122-010-1264-2
[21]	SHAW E, RAJCAN I . Molecular mapping of soybean seed tocopherols in the cross ‘OAC Bayfield’×‘OAC Shire’. Plant Breeding. 2017,136:83-93. doi: 10.1111/pbr.2017.136.issue-1
[22]	李海燕 . 大豆维生素E含量的遗传分析及QTL定位[D]. 哈尔滨: 东北林业大学, 2010.
	LI H Y . Genetic and QTL analysis of the content of vitamin E in soybean[D]. Harbin: Northeast Forestry University, 2010. ( in Chinese)
[23]	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. doi: 10.3724/SP.J.1006.2015.00187
	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. (in Chinese) doi: 10.3724/SP.J.1006.2015.00187
[24]	曹锡文, 刘兵, 章元明 . 植物数量性状分离分析Windows 软件包SEA的研制. 南京农业大学学报, 2013,36(6):1-6.
	CAO X W, LIU B, ZHANG Y M . SEA: A software package of segregation analysis of quantitative traits in plants. Journal of Nanjing Agricultural University, 2013,36(6):1-6. (in Chinese)
[25]	WANG S C, BASTEN C J, ZENG Z B . Windows QTL cartographer 2.5 user manual. Department of Statistics[J/OL], North Carolina State University, Raleigh, NC, 2005.
[26]	王珍 . 大豆SSR遗传图谱构建及重要农艺性状QTL分析[D]. 南宁: 广西大学, 2004.
	WANG Z . Construction of soybean SSR based map and QTL analysis important agronomic traits[D]. Nanning: Guangxi University, 2004. ( in Chinese)
[27]	梁慧珍 . 大豆子粒性状的遗传及QTL分析[D]. 西安: 西北农林科技大学, 2006.
	LIANG H Z . Genetic analysis and QTL mapping of seed traits in soybean( Glycine max, 2006. ( in Chinese)
[28]	SONG Q J, MAREK L F, SHOEMAKER R C, LARK K G, CONCIBIDO V C, DELANNAY X, PECHT J E, CREGAN P B . A new integrated genetic linkage map of the soybean. Theoretical and Applied Genetics, 2004,109:122-128. doi: 10.1007/s00122-004-1602-3
[29]	MCCOUCH S R, CHO Y G, YANO M, PAUL E, BLINSTRUB M, MORISHIMA H, KINOSHITA T . Report on QTL nomenclature. Rice Genetics Newsletter, 1997,14:11-14.
[30]	王金社, 李海旺, 赵团结, 盖钧镒 . 重组自交家系群体4对主基因加多基因混合遗传模型分离分析方法的建立. 作物学报, 2010,36(2):191-201. doi: 10.3724/SP.J.1006.2010.00191
	WANG J S, LI H W, ZHAO T J, GAI J Y . Establishment of segregation analysis of mixed inheritance model with four major genes plus polygenes in recombinant inbred lines population. Acta Agronomica Sinica, 2010,36(2):191-201. (in Chinese) doi: 10.3724/SP.J.1006.2010.00191
[31]	JANSEN R C, VAN OOIJIEN J M, STAM P, DEAN C, JANSEN R C, . Genotype-by-environment interaction in genetic mapping of multiple quantitative trait loci Theoretical and Applied Genetics, 1995,91:33-37. doi: 10.1007/BF00220855
[32]	HAGIWARA W E, ONISH K, TAKAMURE I, SANO Y . Transgressive segregation due to linked QTLs for grain characteristics of rice. Euphytica, 2006,150:27-35. doi: 10.1007/s10681-006-9085-8
[33]	CHANG W, DONG L, WANG Z, HU H, HAN Y, TENG W, ZHANG H, GUO M, LI W . QTL underlying resistance to two HG types of Heterodera glycines found in soybean cultivar ‘L-10’. BMC Genomics, 2011,12:233.
[34]	KAZI S, SHULTZ J, AFZAL J, HASHMI R, JASIM M, BOND J, ARELLI P, LIGHTFOOT D . Iso-lines and inbred-lines confirmed loci that underlie resistance from cultivar ‘Hartwig’ to three soybean cyst nematode populations. Theoretical and Applied Genetics, 2018,131(4):757-773. doi: 10.1007/s00122-018-3063-0
[35]	梁慧珍, 余永亮, 杨红旗, 许兰杰, 董薇, 牛永光, 张海洋, 刘学义, 方宣钧 . 大豆异黄酮及其组分含量的遗传分析与QTL检测. 作物学报, 2015,41(9):1372-1383. doi: 10.3724/SP.J.1006.2015.01372
	LIANG H Z, YU Y L, YANG H Q, XU L J, DONG W, NIU Y G, ZHANG H Y, LIU X Y, FANG X J . Genetic analysis and QTL mapping of isoflavone contents and its components in soybean. Acta Agronomica Sinica, 2015,41(9):1372-1383. (in Chinese) doi: 10.3724/SP.J.1006.2015.01372
[36]	PRIOLLI R H G, CAMPOS J B, STABELLINI N S, PINHEIRO J B, VELLO N A . Association mapping of oil content and fatty acid components in soybean. Euphytica, 2015,203(1):83-96. doi: 10.1007/s10681-014-1264-4
[37]	ZHANG D, SONG H N, CHENG H, HAO D R, WANG H, KAN G Z, JIN H X, YU D Y . The acid phosphatase-encoding gene GmACP1 contributes to soybean tolerance to low phosphorus stress. PLoS Genet, 2014,10(1):e1004061. doi: 10.1371/journal.pgen.1004061
[38]	梁慧珍, 余永亮, 杨红旗, 张海洋, 董薇, 李彩云, 杜华, 巩彭涛, 刘学义, 方宣钧 . 大豆产量及主要农艺性状QTL的上位性互作和环境互作分析. 作物学报, 2014,40(1):37-44. doi: 10.3724/SP.J.1006.2014.00037
	LIANG H Z, YU Y L, YANG H Q, ZHANG H Y, DONG W, LI C Y, DU H, GONG P T, LIU X Y, FANG X J . Epistatic effects and QTL × environment interaction effects of QTLs for yield and agronomic traits in soybean. Acta Agronomica Sinica, 2014,40(1):37-44. (in Chinese) doi: 10.3724/SP.J.1006.2014.00037
[39]	CREGAN P B, JARVIK T, BUSH A L, SHOEMAKER R C, LARK K G, KAHLER A L, KAYA N, VANTOAI T T, LOHNES D G, CHUNG J, SPECHT J E . An integrated genetic linkage map of the soybean genome. Crop Science, 1999,39(5):1464-1490. doi: 10.2135/cropsci1999.3951464x
[40]	梁慧珍, 董薇, 许兰杰, 余永亮, 杨红旗, 谭政委, 许阳, 陈鑫伟 . 不同氮磷钾处理大豆苗期主根长和侧根数的QTL定位分析. 中国农业科学, 2017,50(18):3450-3460. doi: 10.3864/j.issn.0578-1752.2017.18.002
	LIANG H Z, DONG W, XU L J, YU Y L, YANG H Q, TAN Z W, XU Y, CHEN X W . QTL mapping for main root length and lateral root number in soybean at the seedling stage in different N, P and K environments. Scientia Agricultura Sinica, 2017,50(18):3450-3460. (in Chinese) doi: 10.3864/j.issn.0578-1752.2017.18.002
[41]	梁慧珍, 余永亮, 许兰杰, 杨红旗, 董薇, 谭政伟, 李磊, 裴新涌, 刘新梅 . 大豆α-生育酚的遗传与QTL分析. 中国农业科学, 2019,52(1):11-20. doi: 10.3864/j.issn.0578-1752.2019.01.002
	LIANG H Z, YU Y L, XU L J, YANG H Q, DONG W, TAN Z W, LI L, PEI X Y, LIU X M . Inheritance and QTL mapping for α-tocopherol in soybean. Scientia Agricultura Sinica, 2019,52(1):11-20. (in Chinese) doi: 10.3864/j.issn.0578-1752.2019.01.002
[42]	李婕, 王忠, 李永慈, 盖钧镒, 黄中文, 邬荣领 . 异速生长的QTL定位模型及一因多效性扩展. 南京林业大学学报(自然科学版), 2014,38(3):35-39.
	LI J, WANG Z, LI Y C, GAI J Y, HUANG Z W, WU R L . A novel QTL mapping model for allometric growth and pleiotropic extension. Journal of Nanjing Forestry University (Natural Science Edition), 2014,38(3):35-39. (in Chinese)
[43]	HAGIWARA W E, ONISH K, TAKAMURE I, SANO Y . Transgressive segregation due to linked QTLs for grain characteristics of rice. Euphytica, 2006,150:27-35. doi: 10.1007/s10681-006-9085-8
[44]	ZHANG Z H, YU S B, YU T, HUANG Z, ZHU Y G . Mapping quantitative trait loci (QTLs) for seedling-vigor using recombinant inbred lines of rice (Oryza sativa L.). Field Crops Research, 2005,91:161-170. doi: 10.1016/j.fcr.2004.06.004
[45]	方宣钧, 吴为人, 唐纪良 . 作物DNA标记辅助育种. 北京: 科学技术出版社, 2001.
	FANG X J, WU W R, TANG J L. Molecular Marker Assistant Breeding in Crop. Beijing: Science and Technology Press, 2001. ( in Chinese)
[46]	SONG K, SLOCUM M K, OSBORN T C . Molecular marker analysis of genes controlling morphological variation in Brassica rapa (syn. campestris). Theoretical and Applied Genetics, 1995,90:1-10. doi: 10.1007/BF00220989
[47]	HOFIUS D, SONNEWALD U . Vitamin E biosynthesis: Biochemistry meets cell biology. Trends in Plant Science, 2003,8(1):6-8.
[48]	KOCH M, LEMKE R, HEISE K P, MOCK H P . Characterization of γ-tocopherol methyltransferases from Capsicum annuum L. and Arabidopsis thaliana. European Journal of Biochemistry, 2003,270(1):84-92. doi: 10.1046/j.1432-1033.2003.03364.x
[49]	涂世伟 . 甘蓝型油菜γ-生育酚甲基转移酶基因(BnVET4)克隆及其遗传转化的研究[D]. 金华: 浙江师范大学, 2006.
	TU S W . Manipulation of BnVET4 encoding γ-tocopherol methyltransferases in oilseed rape (Brassica napus L.)[D]. Jinhua: Zhejiang Normal University, 2006. ( in Chinese)

年份 Year	平均值Mean (μg·g^-1)		亲本差 P₂—P₁ (μg·g^-1)		RIL变幅RIL range (μg·g^-1)		GCV (%)		遗传率 h²
年份 Year	原阳 Yuanyang	三亚 Sanya	原阳 Yuanyang	三亚 Sanya	原阳 Yuanyang	三亚 Sanya	原阳 Yuanyang	三亚 Sanya	遗传率 h²
2011	111.61	100.83	-51.12	-48.77	25.59—164.15	22.85—158.31	22.66	20.08	0.509
2012	109.92	101.18	-49.90	-51.23	33.65—152.19	27.65—161.02	20.07	16.21	0.437
2015	110.26	100.66	-41.10	-44.26	20.01—158.03	31.03—165.06	24.61	22.12	0.498
平均Mean	110.60	100.88	-48.43	-48.09	20.01—164.15	22.85—165.06	22.59	19.65	0.488

性状 Trait	变异来源 Variation
	基因型Genotypes		环境Environment		基因型×环境 Genotypes×Environment		误差Error
	MS	F	MS	F	MS	F	MS
γ-T	14.1624	2.6972*	0.0358	0.0068	16.6445	3.1698**	5.2509

环境 Environment	原阳2011 Yuanyang2011	原阳2012 Yuanyang2012	原阳2015 Yuanyang2015	三亚2011 Sanya2011	三亚2012 Sanya2012	三亚2015 Sanya2015
原阳2011 Yuanyang2011	1
原阳2012 Yuanyang2012	-0.050	1
原阳2015 Yuanyang2015	0.137	0.043	1
三亚2011 Sanya2011	0.417*	0.023	-0.006	1
三亚2012 Sanya2012	0.307	0.542**	0.303	0.238	1
三亚2015 Sanya2015	0.307	-0.243	0.047	0.356	0.361	1

参数 Parameter		γ-生育酚γ-T		参数 Parameter		γ-生育酚 γ-T
参数 Parameter		原阳Yuanyang	三亚Sanya	参数 Parameter		原阳Yuanyang	三亚Sanya
最适模型Optimal model		MX2-Duplicate-A				MX2-Duplicate-A
一阶参数1st order parameter				二阶参数2nd order parameter
M	2011	8.6329	11.3909	σ_p²	2011	6.3918	2.8311
	2012	10.3476	10.4479		2012	3.8596	3.2818
	2015	7.0225	12.4529		2015	9.3853	3.0353
iab(i^*)	2011	1.6711	0.4329	σ_mg²	2011	3.4311	0.3695
	2012	0.9646	0.5169		2012	1.5250	0.4158
	2015	1.8455	0.4010		2015	5.5337	0.3420
[d]	2011		0.1797	σ_pg²	2011		2.3360
	2012		0.2146		2012		2.7899
	2015		0.1921		2015		2.6270
				h²_mg(%)	2011	53.68	13.05
					2012	39.51	12.67
					2015	58.96	11.27
				h_pg² (%)	2011		82.51
					2012		84.99
					2015		86.55

年份 Year	地点 Location	QTL	染色体 Chr.	标记区间 Marker interval	位置 Position (cM)	置信区间 Confidence interval	LOD	加性效应 Additive	R² (%)
2011	原阳Yuanyang	qγ-A1-1	A1(5)	Satt276—Satt364	12.01	7.01—14.35	3.71	0.622	29.54
		qγ-G-1	G(18)	Satt275—Satt038	0.01	0.00—5.15	2.50	-0.590	8.97
	三亚Sanya	qγ-A2-1	A2(8)	Satt315—Satt632	0.01	0.00—4.78	2.54	-1.235	9.45
		qγ-A1-2	A1(5)	Satt364—Satt382	22.24	18.35—27.11	2.63	0.621	29.55
2012	原阳Yuanyang	qγ-K_2-1	K_2(9)	Satt264—Sat_044	37.68	35.40—37.70	3.14	-1.448	12.15
		qγ-A1-3	A1(5)	Satt382—Satt593	55.77	48.12—56.69	3.18	-0.212	14.20
	三亚Sanya	qγ-A2-2	A2(8)	Satt315—Satt632	2.01	0.67—6.17	2.73	-0.711	14.31
		qγ-G-1	G(18)	Satt275—Satt038	0.01	0.00—4.73	2.56	-0.535	8.12
		qγ-M-1	M(7)	Satt220—Satt323	75.57	73.27—75.60	2.73	-0.658	10.04
2015	原阳Yuanyang	qγ-C1-1	C1(4)	Satt565—Satt578	18.82	18.80—28.97	2.99	0.355	7.29
		qγ-A1-1	A1(5)	Satt276—Satt364	12.01	8.38—15.03	3.81	0.613	28.23
	三亚Sanya	qγ-G-1	G(18)	Satt275—Satt038	0.01	0.00—7.02	2.60	-0.801	7.91