结球甘蓝遗传图谱的构建及抗黑腐病QTL定位

doi:10.7668/hbnxb.20193907

摘要/Abstract

摘要：

为了进一步明确结球甘蓝抗黑腐病性状的遗传基础,选育优质抗病品种,以结球甘蓝高抗黑腐病1号生理小种材料4674为父本,高感材料4673为母本,杂交获得F₁,F₁自交获得152个单株的F₂群体。采用苗期喷雾法对F₂群体进行接病,12~14 d后,按照甘蓝苗期鉴定方法对F₂群体进行表型鉴定。从404个分子标记中筛选得到175个多态性好且条带清晰的标记。随后,使用这175个分子标记对F₂群体进行基因型检测和遗传连锁图谱的构建。最后,结合抗病性表型鉴定数据和遗传图谱对甘蓝抗黑腐病性状的QTL进行标记定位。结果表明:有154对分子标记连锁到9条染色体上,其中包括110对InDel标记和44对SSR标记,覆盖长度714.29 cM,标记间平均图距4.64 cM。共定位到7个QTL位点,其中有3个为主效位点,分别是qBR-7-2、qBR-7-3和qBR-4-3,位于7号染色体的CG842110~CG842482及M29~M39标记间和4号染色体上的CD838151~BOE417,LOD值分别为5.75,3.20,3.47,可解释的表型贡献率分别为16.0%,9.2%,10.0%。

关键词: 结球甘蓝, 黑腐病, F₂群体, 遗传连锁图谱, QTL定位

Abstract:

In order to further definite the genetic basis of black rot resistance in cabbage,and breed high-quality disease resistant cultivars,F₁ progenies were obtained by crossing the inbred line 4674(high resistance to race 1)as male parent,with the inbred line 4673(high susceptibility to race 1)as female parent.The F₂ population containing 152 individuals were obtained by F₁self-crossing. The F₂ population was inoculated by spraying at seedling stage. After 12-14 days, the phenotype of F₂ population was identified according to the identification method of cabbage seedling stage.A total of 175 markers with good polymorphism and clear bands were selected from 404 pairs.Subsequently, the 175 molecular markers were used for genotyping of F₂ population and constructing a genetic linkage map. Finally, the QTL for black rot resistance of cabbage was mapped with phenotypic data and genetic map.The results showed that there were 154 molecular markers linked to nine chromosomes,including 110 pairs of InDel markers and 44 pairs of SSR markers and covering a length of 714.29 cM,and the average distance between markers was 4.64 cM.Seven QTLs were located,of which three major QTLs were qBR-7-2,qBR-7-3 and qBR-4-3,mapped on the genomic markers(CG842110-CG842482 and M29-M39)of chromosome 7,and on the genomic markers between CD838151 and BOE417 of chromosome 4,respectively.The explainable phenotypic variation of QTLs were 16.0%,9.2% and 10.0%,and their LOD values were 5.75,3.20 and 3.47,respectively.

Key words: Cabbage, Black rot, F₂ group, Genetic linkage map, QTL mapping

乐祥庆, 钟雄辉, 崔建, 韩睿, 宋旭朦, 颉建明, 康俊根. 结球甘蓝遗传图谱的构建及抗黑腐病QTL定位[J]. 华北农学报, 2023, 38(3): 167-175. doi: 10.7668/hbnxb.20193907.

YUE Xiangqing, ZHONG Xionghui, CUI Jian, HAN Rui, SONG Xumeng, XIE Jianming, KANG Jungen. Construction of a Genetic Linkage Map and QTL Mapping for Black Rot Resistance in Cabbage[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(3): 167-175. doi: 10.7668/hbnxb.20193907.

http://www.hbnxb.net/CN/Y2023/V38/I3/167

图/表 6

参考文献 39

[1]	方智远. 我国甘蓝产销变化与育种对策[J]. 中国蔬菜, 2008(1): 1-2.
	Fang Z Y. Changes in production and marketing of cabbage and its breeding counter measures in China[J]. China Vegetables, 2008(1): 1-2.
[2]	邢苗苗, 刘星, 孔枞枞, 杨丽梅, 庄木, 张扬勇, 王勇, 方智远, 吕红豪. 甘蓝NLR家族全基因组鉴定、进化分析及在不同病害胁迫下的表达分析[J]. 园艺学报, 2019, 46(4): 723-737. doi:10.16420/j.issn.0513-353x.2018-0804. doi: 10.16420/j.issn.0513-353x.2018-0804
	Xing M M, Liu X, Kong C C, Yang L M, Zhuang M, Zhang Y Y, Wang Y, Fang Z Y, Lü H H. Whole-genome identification and evolutionary analysis of cabbage NLR family genes and their expression profiles in response to various disease stress[J]. Acta Horticulturae Sinica, 2019, 46(4): 723-737. doi: 10.16420/j.issn.0513-353x.2018-0804
[3]	杨丽梅, 方智远. 中国甘蓝遗传育种研究60年[J]. 园艺学报, 2022, 49(10): 2075-2098. doi:10.16420/j.issn.0513-353x.2022-0711. doi: 10.16420/j.issn.0513-353x.2022-0711
	Yang L M, Fang Z Y. Researches on cabbage genetics and breeding in China for 60 years[J]. Acta Horticulturae Sinica, 2022, 49(10): 2075-2098. doi: 10.16420/j.issn.0513-353x.2022-0711
[4]	陈学东. 乌盟地区甘蓝黑腐病的初步调查[J]. 内蒙古农业科技, 1987(1):38-39.
	Chen X D. Preliminary investigation on black rot of cabbage in Urumqi League[J]. Inner Mongolia Agricultural Science and Technology, 1987(1): 38-39.
[5]	Vicente J G, Holub E B. Xanthomonas campestris pv. campestris(cause of black rot of crucifers) in the genomic era is still a worldwide threat to brassica crops[J]. Molecular plant pathology, 2013, 14(1):2-18.doi: 10.1111/j.1364-3703.2012.00833.x. doi: 10.1111/j.1364-3703.2012.00833.x pmid: 23051837
[6]	Massomo S M S, Mortensen C N, Mabagala R B. Newman M A, Hockenhull J. Biological control of black rot (Xanthomonas campestris pv.campestris) of cabbage in Tanzania with Bacillus strains[J]. Journal of Phytopathology, 2004, 152(2):98-105. doi:10.1111/j.1439-0434.2003.00808.x. doi: 10.1111/j.1439-0434.2003.00808.x URL
[7]	朱晓炜, 薄天岳, 陈锦秀, 邰翔, 任云英. 结球甘蓝主要农艺性状基因定位的研究进展[J]. 园艺学报, 2017, 44(9): 1729-1737. doi:10.16420/j.issn.0513-353x.2017-0515. doi: 10.16420/j.issn.0513-353x.2017-0515
	Zhu X W, Bo T Y, Chen J X, Tai X, Ren Y Y. Research progress on mapping of genes associated with main agronomic traits of cabbage[J]. Acta Horticulturae Sinica, 2017, 44(9): 1729-1737. doi: 10.16420/j.issn.0513-353x.2017-0515
[8]	罗冉, 吴委林, 张旸, 李玉花. SSR分子标记在作物遗传育种中的应用[J]. 基因组学与应用生物学, 2010, 29(1): 137-143. doi:10.3969/gab.029.000137. doi: 10.3969/gab.029.000137
	Luo R, Wu W L, Zhang Y, Li Y H. SSR marker and its application to crop genetics and breeding[J]. Genomics and Applied Biology, 2010, 29(1): 137-143.
[9]	Camargo L E A. Mapping of quantitative trait loci controlling resistance of Brassica oleracea to Xanthomonas campestris pv. campestris in the field and greenhouse[J]. Phytopathology, 1995, 85(10): 1296-1300. doi: 10.1094/Phyto-85-1296 URL
[10]	Tonu N N, Doullah M A, Shimizu M, Karim M M, Kawanabe T, Fujimoto R, Okazaki K. Comparison of Positions of QTLs Conferring Resistance to Xanthomonas campestris pv.campestris in Brassica oleracea[J]. American Journal of Plant Sciences, 2013, 4(8A):11-20.doi:10.4236/AJPS.2013.48A002. doi: 10.4236/AJPS.2013.48A002
[11]	孔枞枞. 甘蓝黑腐病抗源筛选和抗性遗传分析及QTL定位[D]. 北京: 中国农业科学院, 2019.
	Kong Z Z. Screening of resistance sources to black rot of cabbage, genetic analysis of resistance and QTL mapping[D]. Beijing: Chinese Academy of Agricultural Sciences, 2019.
[12]	Vicente J G, Conway J, Roberts S J, Taylor J D. Identification and origin of Xanthomonas campestris pv. campestris races and related pathovars[J]. Phytopathology, 2001, 91(5): 492-499. doi:10.1094/PHYTO.2001.91.5.492. doi: 10.1094/PHYTO.2001.91.5.492 pmid: 18943594
[13]	翟文慧, 张涛涛, 胡俊, 张凤兰, 严红. 大白菜黑腐病鉴定的湿度试验及其苗期与成株期抗病性的相关分析[J]. 中国蔬菜, 2010(10): 59-63. doi: 10.3969/j.issn.1004-1389.2008.04.048. doi: 10.3969/j.issn.1004-1389.2008.04.048
	Zhai W H, Zhang T T, Hu J, Zhang F L, Yan H. Humidity test of Chinese cabbage black rot identification and correlation analysis of its seedling and adult-plant resistance[J]. China Vegetables, 2010(10): 59-63.
[14]	芦燕. 大白菜黑腐病病原菌鉴定和抗病性鉴定方法研究[D]. 杨凌: 西北农林科技大学, 2008.
	Lu Y. Study on identification of pathogen and identification of resistance to black rot of Chinese cabbage[D]. Yangling: Northwest A&F University, 2008.
[15]	张鑫鑫, 张恩慧, 吴云锋, 许忠民, 蔡美杰, 孙庆国. 甘蓝黑腐病苗期抗性鉴定方法研究[J]. 中国瓜菜, 2021, 34(6):39-46. doi: 10.3969/j.issn.1673-2871.2021.06.006. doi: 10.3969/j.issn.1673-2871.2021.06.006
	Zhang X X, Zhang E H, Wu Y F, Xu Z M, Cai M J, Sun Q G. Study on the method of identification of resistance to black rot in cabbage at seedling stage[J]. China Cucurbits and Vegetables, 2021, 34(6):39-46.
[16]	Taylor J D, Conway J, Roberts S J, Astley D, Vicente J G. Sources and origin of resistance to Xanthomonas campestris pv. campestris in Brassica genomes[J]. Phytopathology, 2002, 92(1): 105-111. doi:10.1094/PHYTO.2002.92.1.105. doi: 10.1094/PHYTO.2002.92.1.105 pmid: 18944146
[17]	陈果, 孔枞枞, 马洪雪, 邓四平, 季家磊, 王勇, 庄木, 张扬勇, 杨丽梅, 关慧明, 方智远, 吕红豪. 甘蓝黑腐病田间病害分级标准的建立与抗源筛选[J]. 中国蔬菜, 2022(8): 42-48.doi: 10.19928/j.cnki.1000-6346.2022.0034. doi: 10.19928/j.cnki.1000-6346.2022.0034
	Chen G, Kong C C, Ma H X, Deng S P, Ji J L, Wang Y, Zhuang M, Zhang Y Y, Yang L M, Guan H M, Fang Z Y, Lü H H. Development of field disease grading standard and germplasm screening for black rot in cabbage[J]. China Vegetables, 2022(8): 42-48.
[18]	李强, 揭琴, 刘庆昌, 王欣, 马代夫, 翟红, 王玉萍. 甘薯基因组DNA高效快速提取方法[J]. 分子植物育种, 2007, 5(5): 743-746.doi: 10.3969/j.issn.1672-416X.2007.05.026. doi: 10.3969/j.issn.1672-416X.2007.05.026
	Li Q, Jie Q, Liu Q C, Wang X, Ma D F, Zhai H, Wang Y. An efficient and rapid method for sweetpotato genomic DNA extraction[J]. Molecular Plant Breeding, 2007, 5(5): 743-746.
[19]	朱洪运. 结球甘蓝遗传图谱的构建及主要农艺性状的QTL定位[D]. 兰州: 甘肃农业大学, 2013.
	Zhu H Y. Construction of genetic map of cabbage and QTL mapping of main agronomic characters[D]. Lanzhou: Gansu Agricultural University, 2013.
[20]	朱洪运, 颉建明, 郁继华, 简元才, 康俊根. 结球甘蓝 AFLP、SSR、SRAP 标记高密度遗传图谱构建[J]. 华北农学报, 2013, 28(3): 82-87.doi:10.3969/j.issn.1000-7091.2013.03.016 doi: 10.3969/j.issn.1000-7091.2013.03.016
	Zhu H Y, Xie J M, Yu J H, Jian Y C, Kang J G. Construction of a genetic linkage map in cabbage based on AFLP, SSR and SRAP markers[J]. Acta Agriculturae Boreali-Sinica, 2013, 28(3): 82-87.
[21]	Liu S Y, Liu Y M, Yang X H, Tong C B, David E, Parkin Isobel A P, Zhao M X, Ma J X, Yu J Y, Huang S M, Wang X Y, Wang Jun Y, Lu K, Fang Z Y, Ian B, Yang T J, Hu Q, Wang X F, Yue Z, Li H J, Yang L F, Wu J, Zhou Q, Wang W X, King Graham J, Pires J Chris, Lu C X, Wu Z Y, Perumal S, Wang Z, Guo H, Pan S K, Yang L M, Min J M, Zhang D, Jin D C, Li W S, Harry B, Tu J X, Guan M, Qi C. The Brassica oleracea genome reveals the asymmetrical evolution of polyploid genomes[J]. Nature communications, 2014, 5(1):3930. doi: 10.1038/ncomms4930. doi: 10.1038/ncomms4930
[22]	陈登辉, 李海龙, 田多成, 孙超, 颉建明, 康俊根. 结球甘蓝高密度遗传图谱构建及抽薹时间相关QTL定位[J]. 甘肃农业大学学报, 2018, 53(5):21-28.doi: 10.13432/j.cnki.jgsau.2018.05.004 doi: 10.13432/j.cnki.jgsau.2018.05.004
	Chen D H, Li H L, Tian D C, Sun C, Xie J M, Kang J G. Construction of high genetic linkage map and QTL mapping analysis on bolting in cabbage[J]. Journal of Gansu Agricultural University, 2018, 53(5):21-28.
[23]	刘泽慈. 甘蓝染色体片段替换系的构建和黑腐病抗性QTL定位及候选基因分析[D]. 兰州: 甘肃农业大学, 2019. doi:10.27025/d.cnki.ggsnu.2019.000025. doi: 10.27025/d.cnki.ggsnu.2019.000025
	Liu Z C. Construction of cabbage (Brassica oleracea Var.capitata) chromosome segment substitution lines and black rot resistance mapping and candidate genes analysis[D]. Lanzhou: Gansu Agricultural University, 2019.
[24]	朱洪运, 田多成, 颉建明, 简元才, 康俊根. 结球甘蓝抽薹开花时间性状的 QTL 定位及分析[J]. 华北农学报, 2013, 28(5): 1-5. doi: 10.3969/j.issn.1000-7091.2013.03.016. doi: 10.7668/hbnxb.2013.05.001
	Zhu H Y, Tian D C, Xie J M, Jian Y C, Kang J G. QTL mapping and analysis on bolting and flowering time in cabbage[J]. Acta Agriculturae Boreali-Sinica, 2013, 28(5): 1-5.
[25]	Wang W X, Huang S M, Liu Y M, Fang Z Y, Yang L M, Hua W, Yuan S X, Liu S Y, Sun J F, Zhuang M, Zhang Y Y, Zeng A S. Construction and analysis of a high-density genetic linkage map in cabbage (Brassica oleracea L. var.capitata)[J]. BMC Genomics, 2012, 13: 523. doi:10.1186/1471-2164-13-523. doi: 10.1186/1471-2164-13-523
[26]	陈琛, 庄木, 程斐, 刘玉梅, 杨丽梅, 张扬勇, 方智远. 甘蓝EST-SSR标记的建立[C]// 庆祝中国园艺学会创建80周年暨第11次全国会员代表大会论文摘要集, 2009:104.
	Chen C, Zhuang M, Cheng F, Liu Y M, Yang L M, Zhang Y Y, Fang Z Y. Establishment of EST-SSR Marker for Cabbage[C]// Summary of Papers Celebrating the 80th Anniversary of the Establishment of Horticultural Society of China and the 11th National Congress of Members, 2009:104.
[27]	Lü H H, Yang L M, Kang J G, Wang Q B, Wang X W, Fang Z Y, Liu Y M, Zhuang M, Zhang Y Y, Lin Y, Yang Y H, Xie B Y, Liu B, Liu J S. Development of InDel markers linked to Fusarium wilt resistance in cabbage[J]. Molecular Breeding, 2013, 32(4): 961-967. doi:10.1007/s11032-013-9925-x. doi: 10.1007/s11032-013-9925-x URL
[28]	朱德威, 陈庆富. 普通小麦遗传图谱研究现状与展望[J]. 种子, 2010, 29(3):64-69. doi: 10.3969/j.issn.1001-4705.2010.03.019. doi: 10.3969/j.issn.1001-4705.2010.03.019
	Zhu D W, Chen Q F. Status and prospect of common wheats' genetic maps[J]. Seed, 2010, 29(3):64-69.
[29]	张帆, 万雪琴, 潘光堂. 玉米抗穗粒腐病QTL定位[J]. 作物学报, 2007, 33(3):491-496.doi:10.3321/j.issn:0496-3490.2007.03.021. doi: 10.3321/j.issn:0496-3490.2007.03.021
	Zhang F, Wan X Q, Pan G T. Molecular mapping of QTL for resistance to maize ear rot caused by Fusarium moniliforme[J]. Journal of Crops, 2007, 33(3): 491-496.
[30]	杨俊品. 玉米分子遗传图谱构建及数量性状基因定位[D]. 雅安: 四川农业大学, 2001.
	Yang J P. Construction of the molecular linkage map and QTL mapping of quantitative traits in maize[D]. Yaan: Sichuan Agricultural University, 2001.
[31]	Li H, Hearne S, Bänziger M, Li Z, Wang J. Statistical properties of QTL linkage mapping in biparental genetic populations[J]. Heredity, 2010, 105(3): 257-267. doi:10.1038/hdy.2010.56. doi: 10.1038/hdy.2010.56 pmid: 20461101
[32]	刘臣. 蓖麻遗传图谱构建及株高性状的QTL定位分析[D]. 湛江: 广东海洋大学, 2014. doi: 10.7666/d.D522252. doi: 10.7666/d.D522252
	Liu C. Construction of linkage genetic map and QTL mapping for plant height related traits Inricinus commnunis L.[D]. Zhanjiang: Guangdong Ocean University, 2014.
[33]	屠德鹏. 蒺藜苜蓿SSR标记遗传图谱的构建及部分农艺性状的QTL定位[D]. 扬州: 扬州大学, 2011.doi: 10.7666/d.y2049400. doi: 10.7666/d.y2049400
	Tu D P. Construction of Medicago truncatula genetic map by SSR and QTL location of agronomic traits[D]. Yangzhou: Yangzhou University, 2011.
[34]	Röder M S, Korzun V, Wendehake K, Plaschke J, Tixier M H, Leroy P, Ganal M W. A microsatellite map of wheat[J]. Genetics, 1998, 149(4):2007-2023. doi:10.1093/GENETICS/149.4.2007. doi: 10.1093/genetics/149.4.2007 pmid: 9691054
[35]	缪体云. 结球甘蓝遗传图谱的构建及主要农艺性状的QTL定位[D]. 北京: 中国农业科学院, 2007. doi: 10.7666/d.Y1057097. doi: 10.7666/d.Y1057097
	Miao T Y. Construction of a molecular genetic map and mapping of QTL related to main agronomic traits in cabbage (Brassica oleracea var. Capitata)[D]. Beijing: Chinese Academy of Agricultural Sciences, 2007.
[36]	李梅. 结球甘蓝抽薹开花性状的遗传、QTL定位及生理研究[D]. 北京: 中国农业科学院, 2009.
	Li M. Genetic, QTL mapping and physiological studies on bolting and flowering traits of cabbage[D]. Beijing: Chinese Academy of Agricultural Sciences, 2009.
[37]	Doullah M, Mohsin G M, Ishikawa K, Hori H, Okazaki K. Construction of a linkage map and QTL analysis for black rot resistance in Brassica oleracea L.[J]. International Journal of Natural Sciences, 2011, 1(1): 1-6.doi:10.3329/ijns.v1i1.8591. doi: 10.3329/ijns.v1i1.8591 URL
[38]	Kifuji Y, Hanzawa H, Terasawa Y, Ashutosh, Nishio T. QTL analysis of black rot resistance in cabbage using newly developed EST-SNP markers[J]. Euphytica, 2013, 190(2):289-295. doi:10.1007/s10681-012-0847-1. doi: 10.1007/s10681-012-0847-1 URL
[39]	Lee J, Izzah N K, Jayakodi M, Perumal S, Joh H J, Lee H J, Lee S C, Park J Y, Yang K W, Nou I S, Seo J, Yoo J, Suh Y, Ahn K, Lee J H, Choi G J, Yu Y, Kim H, Yang T J. Genome-wide SNP identification and QTL mapping for black rot resistance in cabbage[J]. BMC Plant Biology, 2015, 15: 32. doi:10.1186/s12870-015-0424-6. doi: 10.1186/s12870-015-0424-6 pmid: 25644124

染色体 Chromosome	长度/cM Length	标记数 Number of Markers	平均图距/cM Average distance	最大空隙/cM Biggest interval	最小空隙/cM Smallest interval	平均标记数/cM Average interval
CO1	65.21	17	3.84	12.2	0.1	0.26
CO2	66.71	10	6.67	11.3	2.4	0.15
CO3	72.90	13	5.61	14.4	2.8	0.18
CO4	100.03	13	7.69	16.0	1.7	0.13
CO5	129.47	16	8.09	19.3	2.1	0.12
CO6	67.62	15	4.51	10.9	1.0	0.22
CO7	83.82	33	2.54	12.5	0.1	0.39
CO8	61.24	17	3.60	9.5	1.0	0.28
CO9	67.31	20	3.37	6.5	0.9	0.30
共计	714.29	154	4.64	12.5		0.26

染色体 Chromosome	长度/cM Length	标记数 Number of Markers	平均图距/cM Average distance	最大空隙/cM Biggest interval	最小空隙/cM Smallest interval	平均标记数/cM Average interval
CO1	65.21	17	3.84	12.2	0.1	0.26
CO2	66.71	10	6.67	11.3	2.4	0.15
CO3	72.90	13	5.61	14.4	2.8	0.18
CO4	100.03	13	7.69	16.0	1.7	0.13
CO5	129.47	16	8.09	19.3	2.1	0.12
CO6	67.62	15	4.51	10.9	1.0	0.22
CO7	83.82	33	2.54	12.5	0.1	0.39
CO8	61.24	17	3.60	9.5	1.0	0.28
CO9	67.31	20	3.37	6.5	0.9	0.30
共计	714.29	154	4.64	12.5		0.26

亲本Parents		F₂群体F₂ population
母本4673 Female parent 4673	父本4674 Male parent 4674	中亲值 Median relative value	变幅 Ranee	标准差 Std Dev	平均值 Mean	偏度 Skewness	峰度 Kurtosis	方差 Variance
7	0	3.5	0~7	2.24	3.23	0.28	-0.98	4.98

亲本Parents		F₂群体F₂ population
母本4673 Female parent 4673	父本4674 Male parent 4674	中亲值 Median relative value	变幅 Ranee	标准差 Std Dev	平均值 Mean	偏度 Skewness	峰度 Kurtosis	方差 Variance
7	0	3.5	0~7	2.24	3.23	0.28	-0.98	4.98

QTL	位置/cM Position	连锁群 Linkage group	置信区间/cM Confidence interval	侧翼标记 Flanking markers	LOD值 LOD value	加性效应 Additive effect	遗传效应 Genetic effect	表型贡献率/% Phenotypic variation explained
qBR-4-1	17.7	4	17.7~27.1	8C0101~CD823181	1.93	0.83	正向	5.7
qBR-4-2	48.6	4	48.6~53.8	CD819660~CD728591	2.44	0.89	正向	7.1
qBR-4-3	65.4	4	65.4~77.4	CD838151~BOE417	3.47	0.92	正向	10.0
qBR-6-1	60.7	6	55.0~60.7	BOE569~BOE761	1.63	-0.54	负向	4.8
qBR-7-1	12.2	7	10.3~12.2	BOSF1395~BOSF2672	2.47	0.45	正向	7.2
qBR-7-2	34.3	7	34.0~34.3	CG842110~CG842482	5.75	0.12	正向	16.0
qBR-7-3	71.3	7	71.3~83.8	M29~M39	3.20	0.59	正向	9.2

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