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作物学报 ›› 2021, Vol. 47 ›› Issue (2): 245-261.doi: 10.3724/SP.J.1006.2021.04099

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

马铃薯资源晚疫病抗性的全基因组关联分析

蒋伟1,2(), 潘哲超1,2, 包丽仙1,2, 周福仙2, 李燕山1,2, 隋启君1,2,*(), 李先平1,2,*()   

  1. 1云南省农业科学院经济作物研究所, 云南昆明 650205
    2云贵高原马铃薯与油菜科学观测实验站, 云南昆明 650200
  • 收稿日期:2020-04-30 接受日期:2020-09-13 出版日期:2021-02-12 网络出版日期:2020-09-28
  • 通讯作者: 隋启君,李先平
  • 作者简介:蒋伟, E-mail: jiangweiyaas@hotmail.com
  • 基金资助:
    国家自然科学基金项目(31760409);国家自然科学基金项目(31460368);云南省农业联合重点项目([2017FG001(-012)]);国际马铃薯中心合作项目TON(14.1432.5-001.00)

Genome-wide association analysis for late blight resistance of potato resources

JIANG Wei1,2(), PAN Zhe-Chao1,2, BAO Li-Xian1,2, ZHOU Fu-Xian2, LI Yan-Shan1,2, SUI Qi-Jun1,2,*(), LI Xian-Ping1,2,*()   

  1. 1Industrial Crops Research Institute, Yunnan Academy of Agricultural Science, Kunming 650205, Yunnan, China
    2Scientific Observing and Experimental Station of Potato and Rapeseed in Yunnan-Guizhou Plateau, Kunming 650200, Yunnan, China
  • Received:2020-04-30 Accepted:2020-09-13 Published:2021-02-12 Published online:2020-09-28
  • Contact: SUI Qi-Jun,LI Xian-Ping
  • Supported by:
    National Natural Science Foundation of China(31760409);National Natural Science Foundation of China(31460368);Yunnan Agricultural Foundation Projects([2017FG001(-012)]);International Potato Center Collaboration Program TON(14.1432.5-001.00)

摘要:

广谱抗性基因的挖掘是马铃薯高抗晚疫病品种选育的基础。本研究以288份国际马铃薯中心筛选的晚疫病抗性群体为试验材料, 经过连续2年田间调查, 计算AUDPC和sAUDPC值, 评估群体晚疫病抗性; 利用SLAF-seq方法进行群体简化基因组测序, 通过对晚疫病抗性表型数据的全基因组关联分析, 挖掘晚疫病抗性相关的遗传位点和候选基因, 为晚疫病抗性品种选育和抗病机理研究提供一定的理论和材料基础。结果表明, 晚疫病抗性在288份材料间存在着广泛的遗传差异; 基于5种分析模型, 共鉴定到82个与晚疫病抗性显著关联的位点; 在关联区间关联到54个已知或可能与晚疫病抗性相关的基因。其中, 23个基因为抗性基因, 包括晚疫病抗性基因R1同源基因、Sw-5同源基因(R8)和Rpi-vnt1以及编码多效性耐药蛋白基因; 5个基因编码MAPK蛋白和WRKY转录因子; 1个基因参与茉莉酸途径; 3个基因与水杨酸途径相关; 6个基因是病程相关的基因; 3个基因参与苯基丙酸类合成途径; 其他与晚疫病抗性相关的基因, 如HMGR基因(2个)、细胞色素P450 (21个)。

关键词: 马铃薯, 晚疫病抗性, 全基因组关联分析, 抗性基因, 国际马铃薯中心

Abstract:

The mining of broad-spectrum resistance genes is the basis of the breeding of potato varieties with high late blight resistance. In this study, 288 samples of late disease resistance selected by the International Potato Center (CIP) for years were planted and were investigated for two consecutive years, which could provide some theoretical and material basis for breeding of late blight resistant varieties and help to study on disease resistance mechanism. The late blight resistance of each sample was evaluated using AUDPC and sAUDPC values. The population simplified genome sequencing was performed by SLAF-seq and carried out by genome-wide association analysis in order to discover the genetic loci and candidate genes related to late blight resistance. There were wide genetic differences in resistance to late blight among 288 samples. Through genome-wide association analysis, based on five analytical models, a total of 82 loci significantly correlated with late blight resistance were identified. Combined with genomic annotation, there were 54 candidate genes known or likely to be associated with late blight resistance in the association interval. Among them, 23 genes were resistant genes, including late blight resistance gene R1 homolog, Sw-5 homolog (R8) and Rpi-vnt1, as well as gene encoding pleiotropic resistance protein. Five genes encoded MAPK protein and WRKY transcription factor; one gene was involved in the JA pathway; three genes were associated with salicylic acid pathway; six genes were related to the course of the disease; three genes were involved in the synthesis of phenylpropionic acid. Other genes related to resistance to late blight included HMGR gene (2) and cytochrome P450 (21).

Key words: potato, late blight resistance, genome-wide association analysis, resistance gene, CIP

表1

自国际马铃薯中心引进的288份马铃薯群体材料"

编号
Code
CIP编号
CIP accession
群体
Population
编号
Code
CIP编号
CIP accession
群体
Population
编号
Code
CIP编号
CIP accession
群体
Population
C001 CIP381381.13 A C106 CIP392740.4 LTVR D095 CIP394900.1 BW
C002 CIP391065.69 B3 C107 CIP392745.7 LTVR D096 CIP395011.2 B3
C003 CIP391583.25 B3 C108 CIP392759.1 LTVR D097 CIP395015.6 B3
C004 CIP392617.54 B3 C109 CIP393613.2 LTVR D098 CIP395017.14 B3
C005 CIP392634.52 B3 C110 CIP393615.6 LTVR D099 CIP395017.227 B3
C006 CIP393073.179 B3 C112 CIP397030.31 LTVR D100 CIP395017.229 B3
C007 CIP393073.197 B3 C113 CIP397035.26 LTVR D101 CIP395017.242 B3
C008 CIP393227.66 B3 C114 CIP302428.20 LTVR D102 CIP395037.107 B3
C009 CIP393228.67 B3 C115 CIP302476.108 LTVR D103 CIP395077.12 B3
C010 CIP393371.164 B3 C116 CIP302499.30 LTVR D105 CIP395096.2 B3
C011 CIP391004.18 B3 C117 CIP304345.102 LTVR D106 CIP395109.29 B3
C012 CIP392657.171 B3 C118 CIP304350.100 LTVR D107 CIP395109.34 B3
C013 CIP393280.64 B3 C119 CIP304350.118 LTVR D108 CIP395111.13 B3
C014 CIP391047.34 B3 C120 CIP304350.95 LTVR D109 CIP395112.19 B3
C015 CIP391058.175 B3 C121 CIP304371.67 LTVR D110 CIP395112.32 B3
C016 CIP393085.5 B3 C122 CIP304383.41 LTVR D111 CIP395112.36 B3
C017 CIP398192.213 B3-HT C123 CIP304383.80 LTVR D112 CIP395112.6 B3
C018 CIP398098.119 B3-HT C124 CIP304387.39 LTVR D113 CIP395123.6 B3
C019 CIP398098.203 B3-HT C125 CIP304405.47 LTVR D114 CIP395169.17 B3
C020 CIP398180.253 B3-HT C127 CIP397077.16 LTVR D117 CIP395448.1 BW
C021 CIP398180.289 B3-HT C128 CIP391919.3 BW D118 CIP396004.225 B3
C024 CIP398193.553 B3-HT C129 CIP391930.1 BW D119 CIP396004.263 B3
C025 CIP398203.509 B3-HT C130 CIP391931.1 BW D121 CIP396008.104 B3
C026 CIP398208.219 B3-HT C131 CIP394906.6 BW D123 CIP396009.258 B3
C027 CIP398208.33 B3-HT C132 CIP395438.1 BW D124 CIP396012.266 B3
C028 CIP398208.58 B3-HT C133 CIP394904.20 BW D126 CIP396023.109 B3
C029 CIP398208.704 B3-HT D001 CIP300055.32 LTVR D127 CIP396026.101 B3
编号
Code
CIP编号
CIP accession
群体
Population
编号
Code
CIP编号
CIP accession
群体
Population
编号
Code
CIP编号
CIP accession
群体
Population
C030 CIP301024.14 B3-LTVR D002 CIP300063.9 LTVR D128 CIP396026.103 B3
C031 CIP301029.18 B3-LTVR D003 CIP300065.4 LTVR D130 CIP396031.108 B3
C032 CIP301040.63 B3-LTVR D005 CIP300135.3 LTVR D131 CIP396031.119 B3
C033 CIP300046.22 LTVR D007 CIP301023.15 B3-LTVR D132 CIP396033.102 B3
C034 CIP300048.12 LTVR D008 CIP301024.95 B3-LTVR D133 CIP396034.103 B3
C035 CIP300054.29 LTVR D009 CIP301026.23 B3-LTVR D134 CIP396034.268 B3
C036 CIP300056.33 LTVR D011 CIP301041.26 B3-LTVR D137 CIP396038.101 B3
C037 CIP300063.4 LTVR D012 CIP301044.36 B3-LTVR D138 CIP396038.105 B3
C038 CIP300066.11 LTVR D013 CIP301045.74 B3-LTVR D139 CIP396038.107 B3
C039 CIP300072.1 LTVR D014 CIP301055.53 B3-LTVR D140 CIP396043.226 B3
C040 CIP300093.14 LTVR D015 CIP301056.54 B3-LTVR D142 CIP396063.1 B3-LTVR
C041 CIP300099.22 LTVR D017 CIP303381.106 LTVR D143 CIP396063.16 B3-LTVR
C042 CIP300101.11 LTVR D018 CIP303381.30 LTVR D144 CIP396180.22 B3-LTVR
C043 CIP379706.27 LTVR D019 CIP304347.6 LTVR D145 CIP396240.2 B3
C044 CIP385499.11 LTVR D021 CIP304350.18 LTVR D147 CIP396241.4 B3
C046 CIP388676.1 LTVR D022 CIP304350.78 LTVR D148 CIP396244.12 B3
C047 CIP388972.22 LTVR D023 CIP304351.109 LTVR D149 CIP396268.1 B3-LTVR
C048 CIP390478.9 LTVR D024 CIP304351.31 LTVR D150 CIP396268.9 B3-LTVR
C049 CIP391207.2 LTVR D025 CIP304366.46 LTVR D151 CIP396269.14 B3-LTVR
C050 CIP391382.18 LTVR D028 CIP304371.58 LTVR D152 CIP396269.16 B3-LTVR
C051 CIP392781.1 LTVR D029 CIP304387.17 LTVR D154 CIP396272.18 B3-LTVR
C052 CIP392797.22 LTVR D030 CIP304387.92 LTVR D156 CIP396272.21 B3-LTVR
C053 CIP392822.3 LTVR D031 CIP304394.56 LTVR D157 CIP396272.37 B3-LTVR
C054 CIP392973.48 LTVR D033 CIP304399.5 LTVR D158 CIP396273.48 B3-LTVR
C056 CIP394034.65 LTVR D035 CIP304406.31 LTVR D160 CIP397006.18 LTVR
C057 CIP394034.7 LTVR D036 CIP374080.5 A D162 CIP397039.53 LTVR
C058 CIP394579.36 LTVR D037 CIP377744.1 A D164 CIP397060.19 LTVR
C059 CIP394600.52 LTVR D040 CIP381178.14 A D165 CIP398098.205 B3-HT
C061 CIP394613.139 LTVR D041 CIP381379.12 A D166 CIP398098.231 B3-HT
C062 CIP394613.32 LTVR D042 CIP381381.9 A D167 CIP398098.570 B3-HT
C063 CIP394614.117 LTVR D043 CIP381403.16 A D168 CIP398098.65 B3-HT
C064 CIP394881.8 LTVR D044 CIP384321.3 A D169 CIP398180.144 B3-HT
C065 CIP395186.6 LTVR D047 CIP387164.4 B3 D170 CIP398190.112 B3-HT
C066 CIP395193.6 LTVR D048 CIP389746.2 B3 D171 CIP398190.200 B3-HT
C067 CIP395195.7 LTVR D050 CIP391011.17 B3 D172 CIP398190.404 B3-HT
C068 CIP395196.4 LTVR D051 CIP391046.14 B3 D173 CIP398190.523 B3-HT
C069 CIP395197.5 LTVR D052 CIP391065.81 B3 D174 CIP398190.530 B3-HT
C070 CIP395432.51 LTVR D053 CIP391580.30 B3 D175 CIP398190.571 B3-HT
C071 CIP395434.1 LTVR D054 CIP391585.179 B3 D176 CIP398190.605 B3-HT
C072 CIP395436.8 LTVR D055 CIP391585.5 B3 D177 CIP398190.615 B3-HT
C076 CIP397012.22 LTVR D057 CIP392025.7 LTVR D178 CIP398190.735 B3-HT
C078 CIP397016.7 LTVR D058 CIP392285.72 BW D179 CIP398192.41 B3-HT
C079 CIP397029.21 LTVR D059 CIP392633.64 B3 D180 CIP398192.553 B3-HT
C080 CIP397036.7 LTVR D060 CIP392634.49 B3 D181 CIP398192.592 B3-HT
C081 CIP397039.51 LTVR D061 CIP392637.10 B3 D182 CIP398193.158 B3-HT
编号
Code
CIP编号
CIP accession
群体
Population
编号
Code
CIP编号
CIP accession
群体
Population
编号
Code
CIP编号
CIP accession
群体
Population
C082 CIP397044.25 LTVR D064 CIP392821.1 LTVR D183 CIP398193.650 B3-HT
C083 CIP397055.2 LTVR D065 CIP393075.54 B3 D185 CIP398208.29 B3-HT
C084 CIP397065.2 LTVR D067 CIP393077.54 B3 D186 CIP398208.505 B3-HT
C085 CIP397067.2 LTVR D068 CIP393079.24 B3 D187 CIP398208.620 B3-HT
C086 CIP397069.5 LTVR D069 CIP393079.4 B3 D188 CIP398208.670 B3-HT
C087 CIP397073.15 LTVR D070 CIP393083.2 B3 D189 CIP399001.44 B1
C088 CIP397078.12 LTVR D071 CIP393084.31 B3 D190 CIP399004.19 B1
C089 CIP397079.26 LTVR D072 CIP393220.54 B3 D192 CIP399049.14 B1
C090 CIP397079.6 LTVR D074 CIP393248.55 B3 D193 CIP399049.16 B1
C091 CIP397098.12 LTVR D075 CIP393280.57 B3 D194 CIP399049.22 B1
C092 CIP397099.6 LTVR D077 CIP393284.39 B3 D195 CIP399053.11 B1
C093 CIP397100.9 LTVR D078 CIP393339.242 B3 D196 CIP399053.15 B1
C094 CIP397196.3 LTVR D079 CIP393349.68 B3 D199 CIP399067.22 B1
C095 CIP397196.8 LTVR D080 CIP393371.157 B3 D201 CIP399072.28 B1
C096 CIP397197.9 LTVR D081 CIP393371.159 B3 D203 CIP399075.32 B1
C097 CIP398014.2 LTVR D082 CIP393371.58 B3 D205 CIP399078.11 B1
C098 CIP388611.22 LTVR D083 CIP393382.44 B3 D206 CIP399079.22 B1
C099 CIP388615.22 LTVR D084 CIP393385.39 B3 D207 CIP399083.4 B1
C100 CIP389468.3 LTVR D085 CIP393385.47 B3 D209 CIP399085.23 B1
C101 CIP390637.1 LTVR D088 CIP393617.1 LTVR D210 CIP399085.30 B1
C102 CIP391180.6 LTVR D089 CIP394223.19 LTVR D212 CIP694474.33 PREBRED
C103 CIP391533.1 LTVR D092 CIP394895.7 BW D214 CIP800048 VARIETY
C104 CIP391724.1 LTVR D093 CIP394898.13 BW D216 CIP800923 VARIETY
C105 CIP392032.2 LTVR D094 CIP394899.5 BW D217 CIP800959 VARIETY

图1

288份群体材料AUDPC和sAUDPC频次分布图及相关性散图(2015-2016年) A: 2015年AUDPC值; B: 2016年AUDPC值; C: 2015年sAUDPC值; D: 2016年sAUDPC值; E: 2015年和2016年AUDPC值相关性散布图; F: 2015年和2016年sAUDPC值相关性散布图。"

图2

AUDPC_mean (A: CMLM; C: EMMAX; E: FASTLMM; G: GLM; I: MLM模型)和sAUDPC_mean (B: CMLM; D: EMMAX; F: FASTLMM; H: GLM; J: MLM模型)不同模型下的QQ图"

表2

6个马铃薯晚疫病抗性性状在不同分析模型和显著水平下的显著关联标记"

编号
Number
SNP位点
SNP locus
染色体
Chr.
不同模型、显著水平下与SNP相关联的性状
Traits correlated with SNP under different analytical models and significant level
1 10774833 3 5d_0.1, 6d_0.1
2 10774840 3 5d_0.1
3 10774841 3 5d_0.1
4 14767856 3 3d, e_0.1, 4d, e_0.1, 6d, e_0.1
5 18132045 3 3d_0.1, 4d_0.1
6 32015964 3 3d_0.1, 4d_0.1, 6d_0.1
7 7144093 4 5d_0.1, 6d_0.1
8 17868575 4 3d_0.1, 4d_0.1
9 31375673 4 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
10 4579130 5 3b, c, d_0.1, 3b, c_0.01, 4b, c, d_0.1, 4b_0.01
11 2241058 6 1a, b, c, d, e_0.1, 2a, b, c, d, e_0.1, 1b, c, d_0.01, 2b, c,d_0.01, 3d_0.1, 4d_0.1, 3d_0.01, 4d_0.01, 5a, b, c, d, e_0.1, 6a, b, c, d, e_0.1, 5d_0.01, 6d_0.01
12 2241060 6 1a, b, c, d, e_0.1, 2a, b, c, d, e_0.1, 1b, c, d_0.01, 2b, c,d_0.01, 3d_0.1, 4d_0.1, 3d_0.01, 4d_0.01, 5a, b, c, d, e_0.1, 6 b, c, d, e_0.1, 5d_0.01, 6d_0.01
13 17610976 7 1d_0.1, 2d_0.1, 5d_0.1, 6d_0.1, 5d_0.01
14 37485429 7 1d_0.1, 2d_0.1, 5d_0.1, 6d_0.1
15 22968094 9 1d_0.1, 2d_0.1, 5d_0.1, 6d_0.1
16 30491004 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
17 30518418 9 1d, e_0.1, 1d_0.01, 2d, e_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d, e_0.1, 5d_0.01, 6d, e_0.1, 6d_0.01
18 30522602 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d, e_0.1, 5d_0.01, 6d, e_0.1, 6d_0.01
19 30542106 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
20 30550212 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
21 30568973 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
22 31001787 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
23 31707832 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
24 32170544 9 1d_0.1, 2d_0.1, 3d_0.1, 4d_0.1, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
25 32382888 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
26 32442672 9 1d_0.1
27 32468577 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3 d, e_0.01, 4a, b, c, d, e_0.1, 4d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
28 32627854 9 1a, b, c, d, e_0.1, 1a, b, c, d_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d_0.01, 3b, c, d_0.1, 3 d_0.01, 4b, c, d_0.1, 4d_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d_0.01, 6a, b, c, d, e_0.1, 6b, c, d_0.01
29 32642867 9 1a, c, d, e_0.1, 2a, c, d, e_0.1, 1d_0.01, 2d_0.01, 3d_0.1, 4d_0.1, 3d_0.01, 4d_0.01, 5a, d, e_0.1, 6a, d, e_0.1, 5a, d_0.01, 6d_0.01
30 32747131 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3 b, c, d, e_0.01, 4b, c, d, e_0.1, 4b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5 b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6b, c, d, e_0.01
31 32749768 9 3d_0.1, 4d_0.1
32 32749770 9 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 6d_0.1
编号
Number
SNP位点
SNP locus
染色体
Chr.
不同模型、显著水平下与SNP相关联的性状
Traits correlated with SNP under different analytical models and significant level
33 32752786 9 1d_0.1, 2d_0.1, 3b, d_0.1, 3d_0.01, 4b, d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
34 32753035 9 1d_0.1, 2d_0.1, 3 d_0.1, 3d_0.01, 4 d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
35 32973114 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
36 33118463 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
37 33145926 9 1a, b, c, d, e_0.1, 1a, b, c, d_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d_0.01, 3b, c, d_0.1, 3 c, d_0.01, 4b, c, d_0.1, 4c, d_0.01, 5a, b, c, d, e_0.1, 5 b, c, d_0.01, 6a, b, c, d, e_0.1, 6b, c, d_0.01
38 33145927 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
39 33148740 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
40 33156262 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
41 33170290 9 1d_0.1, 2d_0.1, 5d_0.1, 6d_0.1
42 33170293 9 1d_0.1, 2d_0.1, 5d_0.1, 6d_0.1
43 33211574 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
44 33222811 9 1c, d_0.1, 1d_0.01, 2c, d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5c, d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
45 33229209 9 1d_0.1, 2d_0.1, 5d_0.1
46 33230719 9 3d_0.1, 4d_0.1, 5d_0.1, 6d_0.1
47 33243331 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
48 33244698 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
49 33283731 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
50 33283949 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
51 33288990 9 1d_0.1, 2d_0.1, 3d_0.1, 4d_0.1, 5d_0.1, 6d_0.1
52 33302511 9 1b, c, d_0.1, 1d_0.01, 2b, c, d_0.1, 2d_0.01, 3b, c, d_0.1, 3d_0.01, 4b, c, d_0.1, 4d_0.01, 5b, c, d_0.1, 5d_0.01, 6b, c, d_0.1, 6d_0.01
53 33451096 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
54 33451119 9 1b, c, d_0.1, 1d_0.01, 2b, c, d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5b, c, d_0.1, 5d_0.01, 6b, c, d_0.1, 6d_0.01
55 33480937 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
56 34878453 9 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 6d_0.1, 6d_0.01
57 34898663 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3b, c, d_0.01, 4a, b, c, d, e_0.1, 4b, c, d_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
58 34898695 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3b, c, d, e_0.1, 3b, c, d_0.01, 4 b, c, d, e_0.1, 4b, c, d_0.01, 5a, b, c, d, e_0.1, 5b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
59 34908783 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5a, b, c, d, e_0.1, 5d_0.01, 6a, b, c, d, e_0.1, 6d_0.01
60 34922341 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3 d_0.1, 4 d_0.1, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
61 34935499 9 1a, b, c, d, e_0.1, 1a, c, d_0.01, 2a, b, c, d, e_0.1, 2a, c, d_0.01, 3a, d_0.1, 3d_0.01, 4a, d_0.1, 4d_0.01, 5a, b, c, d, e_0.1, 5a, c, d_0.01, 6a, b, c, d, e_0.1, 6a, d_0.01
62 34959324 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
编号
Number
SNP位点
SNP locus
染色体
Chr.
不同模型、显著水平下与SNP相关联的性状
Traits correlated with SNP under different analytical models and significant level
63 34964634 9 1d_0.1, 2d_0.1
64 34998000 9 1d_0.1, 2d_0.1, 5d_0.1, 6d_0.1
65 35003777 9 1d_0.1, 2d_0.1, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
66 35060347 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
67 35065836 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
68 35097925 9 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
69 35097926 9 3d_0.1, 4d_0.1, 5d_0.1, 6d_0.1
70 35114336 9 3d_0.1, 3d_0.01, 4d_0.1, 4d_0.01, 5d_0.1, 6d_0.1
71 35117629 9 1a, b, c, d, e_0.1, 1a, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, c, d, e_0.01, 3a, d, e_0.1, 3d_0.01, 4a,b, d, e_0.1, 4d_0.01, 5a, b, c, d, e_0.1, 5a, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, d_0.01
72 35123523 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
73 35123542 9 1a, b, c, d, e_0.1, 1b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3b, c, d_0.01, 4a, b, c, d, e_0.1, 4b, c, d_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
74 35123599 9 3d_0.1, 5d_0.1, 6d_0.1
75 35180852 9 1d_0.1, 2d_0.1, 3d_0.1, 4d_0.1, 5d_0.1, 6d_0.1
76 35212385 9 1d_0.1, 1d_0.01, 2 d_0.1, 2d_0.01, 3b, c, d_0.1, 3b, c, d_0.01, 4b, c, d_0.1, 4b, d_0.01, 5b, c, d_0.1, 5d_0.01, 6b, c, d_0.1, 6d_0.01
77 35212534 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
78 35253233 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, d, e_0.1, 3d_0.01, 4a, d, e_0.1, 4d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, d, e_0.01
79 35254558 9 1a, b, c, d, e_0.1, 1a, b, c, d_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d_0.01, 3a,b, c, d, e_0.1, 3b, c, d_0.01, 4a, b, c, d, e_0.1, 4b, c, d_0.01, 5a, b, c, d, e_0.1, 5b, c, d_0.01, 6a, b, c, d, e_0.1, 6b, c, d_0.01
80 35256410 9 1a, b, c, d, e_0.1, 1a, b, c, d, e_0.01, 2a, b, c, d, e_0.1, 2a, b, c, d, e_0.01, 3a, b, c, d, e_0.1, 3a, b, c, d, e_0.01, 4a, b, c, d, e_0.1, 4a, b, c, d, e_0.01, 5a, b, c, d, e_0.1, 5a, b, c, d, e_0.01, 6a, b, c, d, e_0.1, 6a, b, c, d, e_0.01
81 8842711 11 1d_0.1, 2d_0.1, 3d_0.1, 3d_0.01, 4d_0.1, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01
82 23995788 12 1d_0.1, 1d_0.01, 2d_0.1, 2d_0.01, 3 d_0.1, 4 d_0.1, 5d_0.1, 5d_0.01, 6d_0.1, 6d_0.01

图3

GWAS分析生成的曼哈顿图 A: GLM模型分析AUDPC_mean性状; B: GLM模型分析AUDPC_16性状; C: EMMAX模型分析sAUDPC_16性状; D: GLM模型分析AUDPC_15性状。"

表3

根据基因参与途径或分类筛选的晚疫病抗性候选基因"

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Wu Q Y, Huang K, Liu M Y, Zhou Q, Xiong X Y. Research progress of late blight resistance genes in potato. China Potato J, 2014,28:175-179 (in Chines with English abstract).
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