基于全基因组重测序的大豆分子标记开发及籽粒蛋白质含量QTL定位

doi:10.3864/j.issn.0578-1752.2019.16.001

摘要/Abstract

摘要：

【目的】基于全基因组重测序结果,开发与高蛋白、耐荫、抗倒伏等性状紧密相关的分子标记,同时利用开发的分子标记构建遗传连锁图谱,并对籽粒蛋白质含量进行QTL定位,为后续高蛋白、耐荫、抗倒育种研究提供参考和分子标记资源。【方法】以大面积栽培品种南豆12和地方品种十月黄为亲本,构建F₂分离群体。对亲本材料进行覆盖度约为40×的全基因组重测序,用BWA、GATK及Breakdancer等软件比对,检测亲本材料在全基因组范围内的突变类型,挖掘相关变异基因。结合种子不同发育时期和荫蔽处理获得的转录组数据,结合qRT-PCR对发生突变的储藏蛋白、环境适应相关基因进行表达规律分析。同时,基于重测序数据,挖掘亲本间存在于基因编码区的SNP位点,对其进行酶切位点分析,将SNP标记转化为CAPS或dCAPS标记。此外,搜索亲本间存在的插入/缺失变异位点,在插入/缺失位点两侧高度保守的区域设计引物开发InDel标记。对开发的CAPS标记和InDel标记进行多态性筛选,选取具有多态性的CAPS分子标记和InDel标记,对F₂材料进行基因分型。根据分型结果,利用JoinMap 4.0软件进行遗传连锁图谱的构建。依据构建的遗传图谱,结合近红外分析获得F₂材料的籽粒蛋白质含量数据,使用Windows QTL Cartographer V2.5软件对大豆籽粒蛋白质含量进行QTL分析。【结果】测序结果显示,南豆12大量储藏蛋白、环境适应相关的重要基因或同源基因发生突变。转录组数据分析结果显示部分变异基因呈现不同的表达模式且差异显著,qRT-PCR分析进一步验证了该结果。此外,经检测开发的540个CAPS分子标记中有332个具有酶切多态性,300对InDel引物中有201对引物能扩增出多态性。基于533个多态性分子标记构建了一张包含20个连锁群的遗传连锁图谱,覆盖长度2 973.87 cM,标记间平均遗传距离5.58 cM。利用此图谱对大豆籽粒蛋白质含量进行QTL定位,共检测到QTL位点6个,可解释4.68%—18.25%的表型变异。【结论】基于亲本间的变异位点,共开发了533个多态性分子标记（包含8个基因特异性分子标记）,检测到6个大豆籽粒蛋白质含量QTL位点,其中,主效QTL位点1个（qSPC-6）。

关键词: 大豆, 全基因组重测序, 套作, 高蛋白, 耐荫, 抗倒伏, 分子标记

Abstract:

【Objective】 Based on the results of genome-wide re-sequencing, molecular markers closely related to high protein, shade tolerance, lodging resistance and other traits were developed. At the same time, At the same time, genetic linkage maps were constructed using the developed molecular markers, and seed protein content was mapped by QTL, providing reference and molecular marker resources for subsequent research on high protein, shade tolerance and lodging resistance breeding. 【Method】 A F₂ segregating population derived from the cross of Nandou 12 and Shiyuehuang consists of 672 individuals, and two parents were re-sequenced. With the published genome as a reference, the obtained data were assembled with BWA, and explored for the SNP and InDel by GATK and SV by Breakdancer. Carry out expression pattern analysis toward mutational storage proteins and genes related to environmental adaptation by combining with the transcriptome data obtained from different development stages and shade processing of seeds and qRT-PCR, At the same time, based on the resequencing data, excavate the SNP sites in the gene coding region between the parents, analyze the restriction enzyme cutting site and transform the SNP markers into CAPS or dCAPS markers. In addition, search the insertion/deletion mutation site and design primer development InDel marker in highly conserved regions on both sides of the insertion/deletion site. Perform polymorphism screening on the CAPS markers and InDel markers developed, select the CAPS molecular markers and InDel markers with polymorphism and carry out genotyping toward F₂ materials. Utilize JoinMap 4.0 software to construct the genetic linkage map according to the genotyping result. Obtain the seed protein content data of F2 material according to the genetic map constructed by combining with the near-infrared analysis and use Windows QTL Cartographer V2.5 to carry out QTL analysis toward soybean seed protein content. 【Result】 The results showed that a large number of storage proteins and important genes or homologous genes related to environmental adaptation mutated in Nandou 12. The results of transcriptome data analysis showed that some variant genes showed different expression patterns and significant differences and the results were further validated by qRT-PCR analysis. In addition, 332 of the 540 CAPS molecular markers had polymorphic, and 201 of 300 pairs of InDel primers could amplify polymorphism. A genetic linkage map containing 20 linkage groups was constructed based on polymorphic molecular markers, covering 2973.87 cM with an average genetic distance of 5.58 cM. Using this map to map the seed protein content of soybean, six QTL loci were detected, which could explain 4.68%-18.25% phenotypic variation.【Conclusion】 Based on the variation loci among parents, 533 polymorphic molecular markers (including 8 gene-specific molecular markers) were developed. Six QTL loci were detected for seed protein content in soybean, including one major QTL locus (qSPC-6).

Key words: soybean, whole genome re-sequencing, relay intercropping, high protein, shade tolerant, lodging resistance, molecular marker

王嘉, 曾召琼, 梁建秋, 于晓波, 吴海英, 张明荣. 基于全基因组重测序的大豆分子标记开发及籽粒蛋白质含量QTL定位[J]. 中国农业科学, 2019, 52(16): 2743-2757.

WANG Jia, ZENG ZhaoQiong, LIANG JianQiu, YU XiaoBo, WU HaiYing, ZHANG MingRong. Development New Molecular Markers for Quantitative Trait Locus (QTL) Analysis of the Seed Protein Content Based on Whole Genome Re-Sequencing in Soybean[J]. Scientia Agricultura Sinica, 2019, 52(16): 2743-2757.

图/表 8

表1

图1

表2

图2

图3

表3

表4

图4

参考文献 42

[1]	周新安, 年海, 杨文钰, 韩天富 . 南方间套作大豆生产发展的现状与对策(Ⅰ). 大豆科技, 2010(3):1-2.
	ZHAOU X A, NIAN H, YANG W Y, HAN T F . Current situation and countermeasures of intercropping soybean production in the south(Ⅰ). Soybean Science and Technology, 2010(3):1-2. (in Chinese)
[2]	周新安, 年海, 杨文钰, 韩天富 . 南方间套作大豆生产发展的现状与对策(Ⅱ). 大豆科技, 2010(4):1-3.
	ZHOU X A, NIAN H, YANG W Y, HAN T F . Current situation and countermeasures of intercropping soybean production in the south(Ⅱ). Soybean Science and Technology, 2010(4):1-3. (in Chinese)
[3]	周新安, 年海, 杨文钰, 韩天富 . 南方间套作大豆生产发展的现状与对策(Ⅲ). 大豆科技, 2010(5):1-2.
	ZHOU X A, NIAN H, YANG W Y, HAN T F . Current situation and countermeasures of intercropping soybean production in the south(Ⅲ). Soybean Science and Technology, 2010(5):1-2. (in Chinese)
[4]	刘广才 . 不同间套作系统种间营养竞争的差异性及其机理研究[D]. 兰州: 甘肃农业大学, 2005.
	LIU G C . Difference and its mechanism of interspecific nutrition competition in different intercropping systems[D]. Lanzhou: Gansu Agricultural University, 2005. (in Chinese)
[5]	杨文钰, 雍太文, 任万军, 樊高琼, 牟锦毅, 卢学兰 . 发展套作大豆, 振兴大豆产业. 大豆科学, 2008,27(1):1-7.
	YANG W Y, YONG T W, REN W J, FAN G Q, MOU J Y, LU X L . Develop relay-planting soybean, revitalize soybean industry. Soybean Science, 2008,27(1):1-7. (in Chinese)
[6]	YANG F, HUANG S, GAO R C, LIU W G, YONG T W, WANG X C, WU X L, YANG W Y . Growth of soybean seedlings in relay strip intercropping system in relation to light quantity and red: far-red ratio. Field Crops Research, 2014,155(155):245-253.
[7]	张彦威, 李伟, 张礼凤, 王彩洁, 戴海英, 徐冉 . 基于重测序的大豆新品种齐黄34的全基因组变异挖掘. 中国油料作物学报, 2016,38(2):150-158. doi: 10.7505/j.issn.1007-9084.2016.02.003
	ZHANG Y W, LI W, ZHANG L F, WANG C J, DAI H Y, XU R . Genome-wide variations of soybean cultivar Qihuang 34 by whole genome re-sequencing. Chinese Journal of Oil Crop Sciences, 2016,38(2):150-158. (in Chinese) doi: 10.7505/j.issn.1007-9084.2016.02.003
[8]	LAI J, LI R, XU X, JIN W, XU M, ZHAO H, XIANG Z, SONG W, YING K, ZHANG M, JIAO Y, NI P, ZHANG J, LI D, GUO X, YE K, JIAN M, WANG B, ZHENG H, LIANG H, ZHANG X, WANG S, CHEN S, LI J, FU Y, SPRINGER N M, YANG H, WANG J, DAI J, SCHNABLE P S, WANG J . Genome-wide patterns of genetic variation among elite maize inbred lines. Nature Genetics, 2010,42(11):1027-1030.
[9]	杜龙岗, 王美兴 . 玉米SLAF标记的开发及其在玉米果皮纤维素含量BSA分析中的应用. 中国农业科学, 2018,51(8):1421-1430. doi: 10.3864/j.issn.0578-1752.2018.08.001
	DU L G, WANG M X . SLAF-marker development and its application in BSA analysis of cellulose content in pericarp of maize kernel. Scientia Agricultura Sinica, 2018,51(8):1421-1430. (in Chinese) doi: 10.3864/j.issn.0578-1752.2018.08.001
[10]	CHIA J M, SONG C, BRADBURY P J, COSTICH D, DE LEON N, DOEBLEY J, ELSHIRE R J, GAUT B, GELLER L, GLAUBITZ J C, GORE M, GUILL K E, HOLLAND J, HUFFORD M B, LAI J, LI M, LIU X, LU Y, MCCOMBIE R, NELSON R, POLAND J, PRASANNA B M, PYHAJARVI T, RONG T, SEKHON R S, SUN Q, TENAILLON M I, TIAN F, WANG J, XU X, ZHANG Z, KAEPPLER S M, ROSS-IBARRA J, MCMULLEN M D, BUCKLER E S, ZHANG G, XU Y, WARE D . Maize hap map2 identifies extant variation from a genome in flux. Nature Genetics, 2012,44(7):803-807.
[11]	TAKAGI H, ABE A, YOSHIDA K, KOSUGI S, NATSUME S, MITSUOKA C, UEMURA A, UTSUSHI H, TAMIRU M, TAKUNO S, INNAN H, CANO L M, KAMOUN S, TERAUCHI R . QTL-seq: Rapid mapping of quantitative trait loci in rice by whole genome resequencing of DNA from two bulked populations. The Plant Journal, 2013,74(1):174-183.
[12]	MUKESH J, KANHU C M, RAMA S, ROMIKA K, ROHINI G . Genome wide discovery of DNA polymorphisms in rice cultivars with contrasting drought and salinity stress response and their functional relevance. Plant Biotechnology Journal, 2014,12(2):253-264.
[13]	XU X, LIU X, GE S, JENSEN D J, HU F J, LI X, DONG Y, GUTENKUNST R N, FANG L, HUANG L, LI J X, HE W M, ZHANG G J, ZHENG X M, ZHANG F M, LI Y R, YU C, KRISTIANSEN K, ZHANG X Q, WANG J, WRIGHT M, MCCOUCH S, NIELSEN R, WANG J, WANG W . Resequencing 50 accessions of cultivated and wild rice yields markers for identifying agronomically important genes. Nature Biotechnology, 2012,30(1):105-111.
[14]	BAI H, CAO Y G, QUAN J Z, DONG L, LI Z Y, ZHU Y B, ZHU L H, DONG Z P, LI D Y . Identifying the genome-wide sequence variations and developing new molecular markers for genetics research by re-sequencing a landrace cultivar of foxtail millet. PLoS ONE, 2013,8(9):e73514.
[15]	赵庆英, 张瑞娟, 王瑞良, 高建华, 韩渊怀, 杨致荣, 王兴春 . 基于名优谷子品种晋谷21全基因组重测序的分子标记开发. 作物学报, 2018,44(5):686-696.
	ZHAO Q Y, ZHANG R J, WANG R L, GAO J H, HAN Y H, YANG Z R, WANG X C . Genome-wide identification of molecular markers based on genomic re-sequencing of foxtail millet elite cultivar Jingu 21. Acta Agronomica Sinica, 2018,44(5):686-696. (in Chinese)
[16]	岳晓鹏 . 基于甘蓝型油菜基因组重测序开发InDel标记[D]. 武汉: 华中农业大学, 2014.
	YUE X P . Development of Indel markers based on whole genome resequencing in Brassica napus[D]. Wuhan: Huazhong Agricultural University, 2014. (in Chinese)
[17]	LIN T, ZHU G, ZHANG J, XU X, YU Q, ZHENG Z, ZHANG Z, LUN Y, LI S, WANG X, HUANG Z, LI J, ZHANG C, WANG T, ZHANG Y, WANG A, ZHANG Y, LIN K, LI C, XIONG G, XUE Y, MAZZUCATO A, CAUSSE M, FEI Z, GIOVANNONI J J, CHETELAT R T, ZAMIR D, STÄDLER T, LI J, YE Z, DU Y, HUANG S . Genomic analyses provide insights into the history of tomato breeding. Nature Genetics, 2014,46(11):1220-1226.
[18]	KANG Y J, AHN Y K, KIM K T, JUN T H . Resequencing of Capsicum annuum parental lines (YCM334 and Taean) for the genetic analysis of bacterial wilt resistance. BMC Plant Biology, 2016,16(1):235.
[19]	束永俊, 李勇, 柏锡, 才华, 纪巍, 朱延明 . 基于基因重测序信息的大豆基因靶向CAPS标记开发. 作物学报, 2009,35(11):2015-2021. doi: 10.3724/SP.J.1006.2009.02015
	SHU Y J, LI Y, BAI X, CAI H, JI W, ZHU Y M . Development of soybean gene-driven functional caps markers based on gene resequencing. Acta Agronomica Sinica, 2009,35(11):2015-2021. (in Chinese) doi: 10.3724/SP.J.1006.2009.02015
[20]	SONG X, WEI H, CHENG W, YANG S, ZHAO Y, LI X, LUO D, ZHANG H, FENG X . Development of INDEL markers for genetic mapping based on whole genome resequencing in soybean. G3-Genes Genomes Genetics, 2015,5(12):2793-2799.
[21]	DIERS B W, KEIM P, FEHR W R, SHOEMAKER R C . RFLP analysis of soybean seed protein and oil content. Theoretical and Applied Genetics, 1992,83:608-612.
[22]	魏荷, 王金社, 卢为国 . 大豆子粒蛋白质含量分子遗传研究进展. 中国油料作物学报, 2015,37(3):394-410. doi: 10.7505/j.issn.1007-9084.2015.03.021
	WEI H, WANG J S, LU W G . Molecular genetics advances in soybean seed protein. Chinese Journal of Oil Crop Sciences, 2015,37(3):394-410. (in Chinese) doi: 10.7505/j.issn.1007-9084.2015.03.021
[23]	颜彦, 周加权 . 南豆12不同播期的产量效应初探. 种子科技, 2018,36(1):115-116.
	YAN Y, ZHOU J Q . A preliminary study on the yield effect of different sowing dates of Nandou 12. Seed Technology, 2018,36(1):115-116. (in Chinese)
[24]	罗玲, 于晓波, 万燕, 蒋涛, 杜俊波, 邹俊林, 杨文钰, 刘卫国 . 套作大豆苗期倒伏与茎秆内源赤霉素代谢的关系. 中国农业科学, 2015,48(13):2528-2537. doi: 10.3864/j.issn.0578-1752.2015.13.005
	LUO L, YU X B, WAN Y, JIANG T, DU J B, ZOU J L, YANG W Y, LIU W G . The relationship between lodging and stem endogenous gibberellins metabolism pathway of relay intercropping soybean at seedling stage. Scientia Agricultura Sinica, 2015,48(13):2528-2537. (in Chinese) doi: 10.3864/j.issn.0578-1752.2015.13.005
[25]	张明荣, 吴海英 . 四川间套作大豆生产现状与发展分析. 中国种业, 2009,10:16-18.
	ZHANG M R, WU H Y . Analysis of production status and development of relay cropping soybean in Sichuan. China Seed, 2009,10:16-18. (in Chinese)
[26]	STEWART C N J, VIA L E . A rapid CTAB DNA isolation technique useful for RAPD fingerprinting and other PCR applications. BioTechniques, 1993,14:748-750.
[27]	GRANT D, NELSON R T, CANNON S B, SHOEMAKER R C . SoyBase, the USDA-ARS soybean genetics and genomics database. Nucleic Acids Research, 2010,38(suppl. 1):843-846.
[28]	LI H, DURBIN R . Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics, 2009,25:1754-1760.
[29]	MC K A, HANNA M, BANKS E, SIVACHENKO A, CIBULSKIS K, KERNYTSKY A, GARIMELLA K, ALTSHULER D, GABRIELS, D M, DEPRISTO M A . The genome analysis toolkit: A map reduces frame-work for analyzing next-generation DNA sequencing data. Genome Research, 2010,20(9):1297-1303.
[30]	CHEN K, WALLIS J W, MCLELLAN M D, LARSON D E, KALICKI J M, POHL C S, MCGRATH S D, WENDL M C, ZHANG Q Y, LOCKE D P, SHI X Q, FULTON R S, LEY T J, WILSON R K, DING L, MARDIS E R . BreakDancer: An algorithm for high- resolution mapping of genomic structural variation. Nature Methods, 2009,6(9):677-681.
[31]	GONG W Z, QI P, DU J B, SUN X, WU X L, SONG C, LIU W G, WU Y S, YU X B, YONG T W, WANG X C, YANG F, YAN Y H and YANG W Y . Transcriptome analysis of shade-induced inhibition on leaf size in relay intercropped soybean. PLoS ONE, 2014,9(6):e98465.
[32]	NEFF M M, TURK E, KALISHMAN M . Web-based primer design for single nucleotide polymorphism analysis. Trends in Genetics, 2002,18:613-615.
[33]	LANDER E S, BOTSTEIN D . Mapping mendelian factors underlying quantitative traits using RFLP linkage maps. Genetics, 1989,121:185-199.
[34]	DU R C, KHAN A, QIAO Y G, YI Y Y, WANG L, LIU X Y, LI H Q, WANG J S . Expression pattern of abscisic acid insensitive 3 & IT;(ABI3 & IT;) in:Soybean & IT;(Glycine max)& IT; and its interaction mechanism between storage protein gene promoter. International Journal of Agriculture and Biology, 2018,20(4):833-838.
[35]	刘春, 王显生, 麻浩 . 大豆种子贮藏蛋白遗传改良研究进展. 大豆科学, 2008,27(5):866-873.
	LIU C, WANG X S, MAO H . Genetic improvement on soybean seed storage proteins. Soybean Science, 2008,27(5):866-873. (in Chinese)
[36]	VAN K, MCHALE L K . Meta-analyses of QTLs associated with protein and oil contents and compositions in soybean [Glycine max(L.) Merr.] seed. International Journal of Molecular Sciences, 2017,18(6):1180.
[37]	MCKENDRY A L, MC VETTY P B, VOLDENG H D . Inheritance of seed protein and seed oil content in early maturing soybean. Canadian Journal of Genetics and Cytology, 1985,27(5):603-607.
[38]	KARIKARI B, LI S G, BHAT J A, CAO Y G, KONG J J, YANG J Y, GAI J Y, ZHAO T J . Genome-wide detection of major and epistatic effect QTLs for seed protein and oil content in soybean under multiple environments using high-density Bin map. International Journal of Molecular Sciences, 2019,20(4):979.
[39]	JEONG S C, SAGHAI-MAROOF M A . Detection and genotyping of SNPs tightly linked to two disease resistance loci, Rsv1 and Rsv3, of soybean. Plant Breeding, 2004,123(4):305-310.
[40]	ZENG D L, TIAN Z X, RAO Y C, DONG G J, YANG Y L, HUANG L C, LENG Y J, XU J, SUN C, ZHANG G G, HU J, ZHU L, GAO Z Y, HU X M, GUO L B, XIONG G S, WANG Y H, LI J Y, QIAN Q . Rational design of high-yield and superior-quality rice. Nature Plants, 2017,3:17031.
[41]	ZHANG Y H, LIU M F, HE J B, WANG Y F, XING G N, LI Y, YANG S P, ZHAO T J, GAI J Y . Marker‑assisted breeding for transgressive seed protein content in soybean [Glycine max(L.) Merr.]. Theoretical and Applied Genetics, 2015,128(6):1061-1072.
[42]	向达兵 . 钾对套作大豆的抗倒伏效应与提高产量的机理研究[D]. 成都: 四川农业大学, 2012.
	XIANG D B . Studies on effect and mechanism of potassium on lodging-resistance and yield improvement in relay strip intercropped soybean[D]. Chengdu: Sichuan Agricultural University, 2012. (in Chinese)

基因ID Gene ID	基因名称 Gene name	正向引物 Forward primer (5'-3')	反向引物 Reverse primer (5'-3')
Glyma.18G176100	ABI3	CTCATCATGCAAATCCTACAGC	CTATCTGATGCAACTCGACTCT
Glyma.13G123500	Gy5	CAGATATAGAGCACCCAGAGAC	GGTGTTGAAGGATTGTGCTAAG
Glyma.18G175500	MAG2	CAAATTTGGGGCATGGCTAATA	CTTGAAAATTGTTTCGGTTGCG
Glyma.18G226500	CRA1	CTGCAAGAAGTGGTCGTTTTTA	ATTTCAGTTGTTCCATGTTCCG
Glyma.02g248300	PAT2	AAAGGCGTTGAAGCTGGATATA	AATCTCCTATCTGGATCTCCGA
Glyma.11g112800	BBX21	CACAACAGGTTTCTTCTCACTG	TGTTCTTTCTTCAATGGCAGTG
Glyma.14g062100	COI1	GTACATACACGACTCCAAGGAC	TTTCCCTTCAGGTTTAACGACT
Glyma.05g062700	SAV1	GTGGGTCTCCAATGTTTTGTAC	GCAAATTGTTTCAATGGGTGTG
Glyma.18g043000	KAN	TTGGAGTTCACATTAGGGAGAC	AAGTTTGTCACATGTTACCGTG
Glyma.03g227300	PHYA	CAAAGTCTGATAGAGCAGCAAC	CTCACTGTAAGCGATGACCTTA
	ACT11	CGGTGGTTCTATCTTGGCATC	GTCTTTCGCTTCAATAACCCTA

基因ID Gene ID	基因名称 Gene name	注释 Annotation	基因区域内的SNP数目 Genic SNP	CDS中的SNP SNP in CDS	非同义替换SNP Non-syn SNP
Glyma.18G176100	ABI3	ABI3-like转录因子家族成员,促进种子储藏蛋白的积累 A member of ABI3-like transcription factor family, promoting the accumulation of seed storage proteins	72	31	20
Glyma.13G123500	Gy5	大豆球蛋白基因家族成员之一,编码一个11S球蛋白亚基 Encodes an 11S globulin subunit, a member of the soybean globulin gene family	46	33	18
Glyma.11G112800	BBX21	编码B-box锌指转录因子bbx21,与cop1基因相互作用调节避荫 Encodes a B-box zinc finger transcription factor BBX21, genetically interacts with COP1 to regulate shade avoidance.	30	29	16
Glyma.18G175500	MAG2	参与种子贮藏蛋白从内质网到液泡的运输 Involved in transportation of seed storage proteins from the ER to the vacuole	36	27	14
Glyma.13G335200	OBP3	编码一个主要在根中表达的包含转录因子的核定位Dof域 Encodes a nuclear localized Dof domain containing transcription factor expressed primarily in roots.	35	35	14
Glyma.02G057500	SAV1	编码一个油菜素类固醇生物合成途径的限速酶22α羟化酶 Encodes a 22α hydroxylase whose reaction is a rate-limiting step in brassinosteroid biosynthetic pathway	51	29	12
Glyma.08G171800	GA2OX4	编码作用于赤霉素C19的赤霉素2-氧化酶 Encodes a gibberellin 2-oxidase that acts on C19 gibberellins	88	26	11
Glyma.20G048500	VPS26B	细胞内蛋白转运 Intracellular protein transport	32	25	11
Glyma.18G226500	CRA1	编码一个12S种子储存蛋白 Encodes a 12S seed storage protein	72	26	10
Glyma.14G062100	COI1	避荫 Shade avoidance	28	25	10

性状 Trait	亲本 Parent			F₂群体 F₂ population
性状 Trait	南豆12 Nandou 12	十月黄 Shiyuehuang		范围 Range	均值 Mean	方差 Variance	标准差 SD	变异系数 CV	偏度 Skewness	峰度 Kurtosis
蛋白质含量 Protein contents	50.96	41.35		40.34—51.72	45.37	3.47	1.86	4.10	0.16	-0.09

数量性状座位 QTL	染色体 Chromosome	标记区间 Position (cM)	置信区间 Confidence interval	LOD	加性效应 Additive	贡献率 R²(%)
qSPC-1	Gm06(C2)	34.01	GmIn063—GmCA130	8.09	-0.23	5.39
qSPC-2	Gm07(M)	46.71	GmCA168—GmCA171	5.26	0.22	5.36
qSPC-3	Gm15(E)	26.51	GmIn149—GmIn150	4.53	0.21	4.68
qSPC-4	Gm15(E)	72.21	GmIn152—GmIn163	9.93	-0.29	9.32
qSPC-5	Gm20(I)	35.61	GmCA493—GmIn285	8.40	-0.47	12.04
qSPC-6	Gm20(I)	103.41	GmdCA522—GmCA537	15.06	-0.68	18.86