Genome-Wide Association Study of Powdery Mildew Resistance of Cucumber Core Germplasms

doi:10.16420/j.issn.0513-353x.2021-0205

Acta Horticulturae Sinica ›› 2021, Vol. 48 ›› Issue (7): 1371-1385.doi: 10.16420/j.issn.0513-353x.2021-0205

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Genome-Wide Association Study of Powdery Mildew Resistance of Cucumber Core Germplasms

JIA Huixia¹, LI Xixiang¹, SONG Jiangping¹, LIN Yu’e², ZHANG Xiaohui¹, QIU Yang¹, YANG Wenlong¹, LOU Qunfeng³, WANG Haiping¹^,^*()

¹Institute of Vegetables and Flowers,Chinese Academy of Agricultural Sciences/Key Laboratory of Horticultural Crop Biology and Germplasm Creation in the Ministry of Agriculture,Beijing 100081,China
²Guangdong Key Laboratory for New Technology Research of Vegetables,Vegetable Research Institute,Guangdong Academy of Agricultural Sciences,Guangzhou 510640,China
³State Key Laboratory of Crop Genetics and Germplasm Enhancement,College of Horticulture,Nanjing Agricultural University,Nanjing 210095,China

Received:2021-02-22 Revised:2021-04-06 Online:2021-07-25 Published:2021-08-10
Contact: WANG Haiping E-mail:wanghaiping@caas.cn

Abstract

Abstract:

In this study,231 cucumber core germplasms collected from the world were used for resistance identification of powdery mildew in Beijing and Guangzhou in 2017 and 2018,respectively. A total of 950 382 SNPs detected by genome re-sequencing were used for genome-wide association analysis. The results showed that the natural incidence of cucumber powdery mildew in Guangzhou was significantly higher than that in Beijing. High correlation was detected between the two-year disease index among different germplasms. Based on the mixed linear model,12 significantly associated signals were detected,and they distributed on chromosomes 1,5 and 6. Among them,pm1.1 was detected repeatedly in two-year in Beijing,while pm5.7 was detected repeatedly in two-year in Guangzhou. These significantly associated signals contained 68 SNPs that exceeded the threshold of-log₁₀(P value) with 6.0. Among them,a non-synonymous mutant SNP（Chr5：16 676 542）located on the exon of xenotropic and polytropic retrovirus receptor（Csa5G471600）. A total of 63 candidate genes were found in the LD block of SNPs associated with powdery mildew resistance,among which seven candidate genes showed significant differential expression after being infected with powdery mildew in the cucumber material with high resistance.

Key words: cucumber, powdery mildew, genome-wide association study, single nucleotide polymorphism, candidate gene

CLC Number:

S642.2

JIA Huixia, LI Xixiang, SONG Jiangping, LIN Yu’e, ZHANG Xiaohui, QIU Yang, YANG Wenlong, LOU Qunfeng, WANG Haiping. Genome-Wide Association Study of Powdery Mildew Resistance of Cucumber Core Germplasms[J]. Acta Horticulturae Sinica, 2021, 48(7): 1371-1385.

Figures/Tables 8

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国家 Country	省 Province	样本数 Number of samples	国家 Country	样本数 Number of samples
中国China	广东Guangdong	39	美国America	10
	北京Beijing	14	日本Japan	7
	山东Shandong	14	印度India	6
	辽宁Liaoning	12	德国Germany	5
	云南Yunnan	12	俄罗斯Russia	4
	吉林Jilin	11	土耳其Turkey	3
	河北Hebei	9	匈牙利Hungary	3
	四川Sichuan	7	伊朗Iran	3
	天津Tianjin	7	阿富汗Afghanistan	2
	河南Henan	5	荷兰Holland	2
	湖北Hubei	5	挪威Norway	2
	湖南Hunan	5	西班牙Spain	2
	福建Fujian	3	叙利亚Syria	2
	黑龙江Heilongjiang	3	以色列Israel	2
	内蒙Inner Mongolia	3	埃及Egypt	1
	山西Shanxi	3	巴基斯坦Pakistan	1
	安徽Anhui	2	波多黎各Puerto Rico	1
	贵州Guizhou	2	丹麦Denmark	1
	新疆Xinjiang	2	哈萨克斯坦Kazakhstan	1
	重庆Chongqing	2	津巴布韦Zimbabwe	1
	广西Guangxi	1	克罗地亚Croatia	1
	江苏Jiangsu	1	肯尼亚Kenya	1
	江西Jiangxi	1	毛里求斯Mauritius	1
	昆明Kunming	1	尼泊尔Nepal	1
	宁夏Ningxia	1	塞尔维亚Serbia	1
			泰国Thailand	1
			希腊Greece	1

国家 Country	省 Province	样本数 Number of samples	国家 Country	样本数 Number of samples
中国China	广东Guangdong	39	美国America	10
	北京Beijing	14	日本Japan	7
	山东Shandong	14	印度India	6
	辽宁Liaoning	12	德国Germany	5
	云南Yunnan	12	俄罗斯Russia	4
	吉林Jilin	11	土耳其Turkey	3
	河北Hebei	9	匈牙利Hungary	3
	四川Sichuan	7	伊朗Iran	3
	天津Tianjin	7	阿富汗Afghanistan	2
	河南Henan	5	荷兰Holland	2
	湖北Hubei	5	挪威Norway	2
	湖南Hunan	5	西班牙Spain	2
	福建Fujian	3	叙利亚Syria	2
	黑龙江Heilongjiang	3	以色列Israel	2
	内蒙Inner Mongolia	3	埃及Egypt	1
	山西Shanxi	3	巴基斯坦Pakistan	1
	安徽Anhui	2	波多黎各Puerto Rico	1
	贵州Guizhou	2	丹麦Denmark	1
	新疆Xinjiang	2	哈萨克斯坦Kazakhstan	1
	重庆Chongqing	2	津巴布韦Zimbabwe	1
	广西Guangxi	1	克罗地亚Croatia	1
	江苏Jiangsu	1	肯尼亚Kenya	1
	江西Jiangxi	1	毛里求斯Mauritius	1
	昆明Kunming	1	尼泊尔Nepal	1
	宁夏Ningxia	1	塞尔维亚Serbia	1
			泰国Thailand	1
			希腊Greece	1

染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	P值 P value
染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	2017- Beijing	2018- Beijing	2017- Guangzhou	2018- Guangzhou
Chr1	pm1.1	20 946 043	A/C	基因间区 Intergenic		7.34E-07
		20 946 325	C/T	基因间区 Intergenic		1.72E-08
		20 946 384	C/T	基因间区 Intergenic		6.04E-08
		20 946 425	A/G	基因间区 Intergenic		2.44E-07
		20 946 565	C/T	基因间区 Intergenic		1.67E-08
		20 947 445	T/C	基因间区 Intergenic		2.13E-08
		20 947 746	G/T	基因间区 Intergenic		4.18E-08
		20 947 997	G/A	基因间区 Intergenic		1.53E-08
		20 948 176	T/A	基因间区 Intergenic		4.91E-09
		20 948 443	C/T	基因间区 Intergenic		3.54E-08
		20 949 791	A/G	上游区 Upstream		2.41E-08
		20 950 222	T/C	内含子区 Intronic		1.17E-08
		20 950 313	T/C	内含子区 Intronic	6.97E-07	1.17E-08
		20 951 178	T/C	内含子区 Intronic		9.50E-08
		20 951 719	G/A	内含子区 Intronic		2.86E-08
		20 951 938	G/A	内含子区 Intronic		2.45E-08
		20 952 168	T/C	内含子区 Intronic		1.43E-08
		20 952 590	A/G	内含子区 Intronic	3.64E-08	1.59E-09
		20 952 617	G/A	内含子区 Intronic	8.71E-08	1.24E-09
		20 953 029	T/C	内含子区 Intronic		7.98E-09
		20 953 187	T/C	内含子区 Intronic		9.32E-09
		20 953 655	A/C	内含子区 Intronic		5.23E-09
		20 954 157	T/C	内含子区 Intronic	1.21E-07	3.17E-09
		20 955 127	T/C	内含子区 Intronic		8.56E-08
		20 955 284	T/A	内含子区 Intronic		6.32E-10
		20 955 289	G/A	内含子区 Intronic	3.57E-07	1.25E-09
		20 955 466	A/C	内含子区 Intronic	3.12E-07	8.79E-09
		20 955 556	G/A	内含子区 Intronic	1.78E-07	4.36E-09
		20 956 182	A/C	外显子区（同义突变）Exonic（synonymous）		1.43E-08
		20 956 822	T/A	内含子区 Intronic		1.34E-08
		20 959 137	G/A	下游区 Downstream		7.29E-07
		20 960 732	T/C	内含子区 Intronic		7.54E-08
		20 960 788	C/T	内含子区 Intronic		2.25E-08
		20 961 611	A/C	内含子区 Intronic		1.23E-08
		20 963 321	A/G	上游区/下游区 Upstream/Downstream		2.31E-07
		20 963 487	A/G	上游区/下游区 Upstream/Downstream	5.23E-07	3.40E-09
		20 963 500	T/C	上游区/下游区 Upstream/Downstream		6.56E-08
		20 964 042	A/G	内含子区 Intronic		9.32E-07
		20 964 068	T/C	内含子区 Intronic		9.32E-07
		20 964 073	T/G	内含子区 Intronic		1.79E-07
		20 964 207	A/G	内含子区 Intronic		1.51E-07
		20 965 920	G/C	内含子区 Intronic		1.30E-07
		20 966 914	T/A	上游区 Upstream		3.98E-07
		20 967 021	C/T	上游区 Upstream		8.88E-07
	pm1.2	21 133 564	C/T	基因间区 Intergenic	4.74E-07
	pm1.3	21 453 973	T/C	内含子区 Intronic	6.68E-07
		21 454 062	A/G	内含子区 Intronic	5.04E-07
		21 459 154	T/C	基因间区Intergenic	7.54E-07
		21 459 157	G/A	基因间区Intergenic	2.46E-07
		21 459 889	A/T	上游区 Upstream	1.46E-07
		21 459 973	C/T	上游区 Upstream	1.29E-07
		21 461 960	A/G	内含子区 Intronic	1.74E-07
染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	P值 P value
染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	2017- Beijing	2018- Beijing	2017- Guangzhou	2018- Guangzhou
Chr5	pm5.1	1 858 575	C/T	上游区 Upstream				4.14E-08
	pm5.2	7 910 694	C/T	基因间区 Intergenic	3.20E-07
	pm5.3	15 769 974	G/C	基因间区 Intergenic				9.23E-07
	pm5.4	16 149 984	G/A	内含子区 Intronic				5.83E-07
	pm5.5	16 316 760	T/A	内含子区 Intronic				1.13E-07
	pm5.5	16 327 596	G/C	内含子区 Intronic				6.55E-07
	pm5.6	16 357 208	T/C	内含子区 Intronic				4.66E-07
		16 380 085	A/T	内含子区 Intronic				1.38E-08
		16 390 016	G/A	上游区 Upstream				2.96E-07
	pm5.7	16 666 839	C/G	基因间区 Intergenic			4.64E-07	5.95E-08
		16 676 542	G/C	外显子区（非同义突变Exonic（Nonsynonymous）			2.44E-10	2.70E-07
		16 683 577	C/A	基因间区 Intergenic			2.24E-08	4.22E-11
	pm5.8	22 992 521	A/G	基因间区 Intergenic		3.44E-08
		22 992 567	A/T	基因间区 Intergenic		3.34E-07
		22 993 739	T/A	基因间区 Intergenic		8.32E-07
Chr6	pm6.1	15 844 016	T/G	基因间区 Intergenic				6.97E-07

染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	P值 P value
染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	2017- Beijing	2018- Beijing	2017- Guangzhou	2018- Guangzhou
Chr1	pm1.1	20 946 043	A/C	基因间区 Intergenic		7.34E-07
		20 946 325	C/T	基因间区 Intergenic		1.72E-08
		20 946 384	C/T	基因间区 Intergenic		6.04E-08
		20 946 425	A/G	基因间区 Intergenic		2.44E-07
		20 946 565	C/T	基因间区 Intergenic		1.67E-08
		20 947 445	T/C	基因间区 Intergenic		2.13E-08
		20 947 746	G/T	基因间区 Intergenic		4.18E-08
		20 947 997	G/A	基因间区 Intergenic		1.53E-08
		20 948 176	T/A	基因间区 Intergenic		4.91E-09
		20 948 443	C/T	基因间区 Intergenic		3.54E-08
		20 949 791	A/G	上游区 Upstream		2.41E-08
		20 950 222	T/C	内含子区 Intronic		1.17E-08
		20 950 313	T/C	内含子区 Intronic	6.97E-07	1.17E-08
		20 951 178	T/C	内含子区 Intronic		9.50E-08
		20 951 719	G/A	内含子区 Intronic		2.86E-08
		20 951 938	G/A	内含子区 Intronic		2.45E-08
		20 952 168	T/C	内含子区 Intronic		1.43E-08
		20 952 590	A/G	内含子区 Intronic	3.64E-08	1.59E-09
		20 952 617	G/A	内含子区 Intronic	8.71E-08	1.24E-09
		20 953 029	T/C	内含子区 Intronic		7.98E-09
		20 953 187	T/C	内含子区 Intronic		9.32E-09
		20 953 655	A/C	内含子区 Intronic		5.23E-09
		20 954 157	T/C	内含子区 Intronic	1.21E-07	3.17E-09
		20 955 127	T/C	内含子区 Intronic		8.56E-08
		20 955 284	T/A	内含子区 Intronic		6.32E-10
		20 955 289	G/A	内含子区 Intronic	3.57E-07	1.25E-09
		20 955 466	A/C	内含子区 Intronic	3.12E-07	8.79E-09
		20 955 556	G/A	内含子区 Intronic	1.78E-07	4.36E-09
		20 956 182	A/C	外显子区（同义突变）Exonic（synonymous）		1.43E-08
		20 956 822	T/A	内含子区 Intronic		1.34E-08
		20 959 137	G/A	下游区 Downstream		7.29E-07
		20 960 732	T/C	内含子区 Intronic		7.54E-08
		20 960 788	C/T	内含子区 Intronic		2.25E-08
		20 961 611	A/C	内含子区 Intronic		1.23E-08
		20 963 321	A/G	上游区/下游区 Upstream/Downstream		2.31E-07
		20 963 487	A/G	上游区/下游区 Upstream/Downstream	5.23E-07	3.40E-09
		20 963 500	T/C	上游区/下游区 Upstream/Downstream		6.56E-08
		20 964 042	A/G	内含子区 Intronic		9.32E-07
		20 964 068	T/C	内含子区 Intronic		9.32E-07
		20 964 073	T/G	内含子区 Intronic		1.79E-07
		20 964 207	A/G	内含子区 Intronic		1.51E-07
		20 965 920	G/C	内含子区 Intronic		1.30E-07
		20 966 914	T/A	上游区 Upstream		3.98E-07
		20 967 021	C/T	上游区 Upstream		8.88E-07
	pm1.2	21 133 564	C/T	基因间区 Intergenic	4.74E-07
	pm1.3	21 453 973	T/C	内含子区 Intronic	6.68E-07
		21 454 062	A/G	内含子区 Intronic	5.04E-07
		21 459 154	T/C	基因间区Intergenic	7.54E-07
		21 459 157	G/A	基因间区Intergenic	2.46E-07
		21 459 889	A/T	上游区 Upstream	1.46E-07
		21 459 973	C/T	上游区 Upstream	1.29E-07
		21 461 960	A/G	内含子区 Intronic	1.74E-07
染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	P值 P value
染色体 Chromosome	显著关联信号 Significantly associated signals	SNP位置/bp SNP Position	基因型 Genotype	SNP注释 SNP annotation	2017- Beijing	2018- Beijing	2017- Guangzhou	2018- Guangzhou
Chr5	pm5.1	1 858 575	C/T	上游区 Upstream				4.14E-08
	pm5.2	7 910 694	C/T	基因间区 Intergenic	3.20E-07
	pm5.3	15 769 974	G/C	基因间区 Intergenic				9.23E-07
	pm5.4	16 149 984	G/A	内含子区 Intronic				5.83E-07
	pm5.5	16 316 760	T/A	内含子区 Intronic				1.13E-07
	pm5.5	16 327 596	G/C	内含子区 Intronic				6.55E-07
	pm5.6	16 357 208	T/C	内含子区 Intronic				4.66E-07
		16 380 085	A/T	内含子区 Intronic				1.38E-08
		16 390 016	G/A	上游区 Upstream				2.96E-07
	pm5.7	16 666 839	C/G	基因间区 Intergenic			4.64E-07	5.95E-08
		16 676 542	G/C	外显子区（非同义突变Exonic（Nonsynonymous）			2.44E-10	2.70E-07
		16 683 577	C/A	基因间区 Intergenic			2.24E-08	4.22E-11
	pm5.8	22 992 521	A/G	基因间区 Intergenic		3.44E-08
		22 992 567	A/T	基因间区 Intergenic		3.34E-07
		22 993 739	T/A	基因间区 Intergenic		8.32E-07
Chr6	pm6.1	15 844 016	T/G	基因间区 Intergenic				6.97E-07

显著关联信号 Significantly associated signals	LD区段/bp LD block	候选基因ID Candidate genes ID	功能注释 Functional annotation
pm1.1	Chr1：20 913 000 ~	Csa1G569320	泛素蛋白连接酶RFI2 E3 ubiquitin-protein ligase RFI2
	21 000 000	Csa1G569330	丝氨酸/精氨酸富集剪接因子SR30 Serine/arginine-rich splicing factor SR30
		Csa1G569340	嘌呤渗透酶11 Probable purine permease 11
		Csa1G569350	磷酸葡萄糖胺变位酶 Phosphoglucosamine mutase
		Csa1G569360	缺氧诱导蛋白 Hypoxia induced protein
		Csa1G569370	SOSS复合体亚基B同系物 SOSS complex subunit B homolog
		Csa1G569380	60S核糖体蛋白L22-2 60S Ribosomal protein L22-2
		Csa1G569390	内质网膜蛋白复合体亚基6 ER membrane protein complex subunit 6
		Csa1G569400	休眠相关蛋白3 Dormancy-associated protein 3
		Csa1G569410	Tafazzin蛋白 Tafazzin protein
		Csa1G569420	未知蛋白Unknown protein
		Csa1G569430	橡胶延长因子Rubber elongation factor
		Csa1G569440	重金属相关异戊二烯化植物蛋白 Heavy metal-associated isoprenylated plant protein
		Csa1G569450	呼吸爆发氧化酶同源蛋白B Respiratory burst oxidase homolog protein B
		Csa1G569460	Formin蛋白11 Formin protein 11
pm1.2	Chr1：21 130 000 ~	Csa1G570200	短链脱氢酶/还原酶 Short-chain dehydrogenase/reductase
	21 137 000	Csa1G570210	核糖体生物发生蛋白类BRX1蛋白 Ribosome biogenesis protein BRX1-like protein
pm1.3	Chr1：21 452 000 ~	Csa1G573670	不依赖sec移位酶TATA Sec-independent translocase protein TATA
	21 468 000	Csa1G573680	转录因子 bHLH transcription factor bHLH（UNE12）
pm5.1	Chr5：1 800 000 ~	Csa5G055570	未知蛋白 Unknown protein
	1 886 000	Csa5G056070	未知蛋白 Unknown protein
		Csa5G056080	富含羟脯氨酸糖蛋白Hydroxyproline-rich glycoprotein-like protein
		Csa5G056090	富含羟脯氨酸糖蛋白Hydroxyproline-rich glycoprotein-like protein
		Csa5G056100	邻氨基苯甲酸盐合酶β亚单元Anthranilate synthase beta subunit 2
		Csa5G056110	未知蛋白 Unknown protein
		Csa5G056610	未知蛋白 Unknown protein
		Csa5G056620	3-（3-羟苯基）丙酸盐/3-羟基桂皮酸羟化酶3-（3-hydroxy-phenyl）propionate/3-hydroxycinnamic acid hydroxylase
		Csa5G056630	香豆素酰基辅酶a：花青素3-O-葡萄糖-6"-O-香豆素酰基转移酶 2 Coumaroyl-CoA：anthocyanidin 3-O-glucoside-6''-O-coumaroyltransferase 2
pm5.2	Chr5：7 910 000 ~	Csa5G180870	RNA结合蛋白 RNA binding protein
	7 921 000	Csa5G180880	三角状五肽重复蛋白Pentatricopeptide repeat-containing protein
pm5.3	Chr5：15 700 000 ~	Csa5G431540	花期控制蛋白 Flowering time control protein FY
	15 847 000	Csa5G432540	泛素羧基末端水解酶 Ubiquitin carboxyl-terminal hydrolase
		Csa5G432550	泛素羧基末端水解酶 Ubiquitin carboxyl-terminal hydrolase
		Csa5G434550	双因子响应调节蛋白 Two-component response regulator ORR10
		Csa5G435050	DNA定向RNA聚合酶V亚单位1 DNA-directed RNA polymerase V subunit 1
		Csa5G435060	未知蛋白 Unknown protein
		Csa5G435070	磷脂酰肌醇4-磷酸5-激酶6 Phosphatidylinositol 4-phosphate 5-kinase 6
		Csa5G435080	醌氧化还原酶NAD（P）H：quinone oxidoreductase
pm5.4	Chr5：16 130 000 ~	Csa5G456730	尿苷5'-磷酸合酶 Uridine 5'-monophosphate synthase
	16 170 000	Csa5G457230	ADP核糖基化因子GTP激活蛋白AGD11 同类型X1蛋白 ADP-ribosylation factor GTPase-activating protein AGD11 isoform X1
		Csa5G457240	对红光的敏感性降低蛋白 Sensitivity to red light reduced protein
pm5.5	Chr5：16 312 000 ~	Csa5G466330	rRNA-加工蛋白质UTP23 rRNA-processing protein UTP23
	16 354 000	Csa5G466340	未知蛋白Unknown protein
		Csa5G466350	富含脯氨酸受体蛋白激酶PERK10 Proline-rich receptor-like protein kinase PERK10
		Csa5G466360	L型凝集素结构域的受体激酶L-type lectin-domain containing receptor kinase
pm5.6	Chr5：16 355 000 ~ 16 382 000	Csa5G466370	组氨酸生物合成双功能蛋白hisIE Histidine biosynthesis bifunctional protein hisIE
		Csa5G466870	40S核糖体蛋白S27 40S ribosomal protein S27
显著关联信号 Significantly associated signals	LD区段/bp LD block	候选基因ID Candidate genes ID	功能注释 Functional annotation
pm5.7	Chr5：16 665 000 ~ 16 706 000	Csa5G471600	异嗜性和多嗜性逆转录病毒受体蛋白 Xenotropic and polytropic retrovirus receptor protein
		Csa5G472100	肽基脯氨酸顺反式异构酶NIMA相互作用蛋白4 Peptidyl-prolyl cis-trans isomerase NIMA-interacting 4
pm5.8	Chr5：22 994 000 ~	Csa5G606690	三角状五肽重复蛋白 Pentatricopeptide repeat-containing protein
	23 053 000	Csa5G606700	未知蛋白 Unknown protein
		Csa5G606710	未知蛋白 Unknown protein
		Csa5G606720	未知蛋白 Unknown protein
		Csa5G606730	Ninja家族蛋白 Ninja-family protein AFP3
		Csa5G606740	天冬酰胺合成酶蛋白 Asparagine synthetase domain-containing protein 1
		Csa5G606750	细胞分裂控制蛋白48同源物B Cell division control protein 48 homolog B
		Csa5G606760	核糖体成熟因子rimP Ribosome maturation factor rimP
		Csa5G606770	6-磷酸葡萄糖酸内酯酶1 6-phosphogluconolactonase 1
		Csa5G606780	CRABS CLAW蛋白CRABS CLAW protein
		Csa5G606790	类大谷粒蛋白EBIG GRAIN 1-like E
pm6.1	Chr6：15 801 000 ~	Csa6G351870	细胞壁连接的类受体激酶3 Wall-associated receptor kinase 3
	15 847 000	Csa6G352870	细胞壁连接的类受体激酶2 Wall-associated receptor kinase 2
		Csa6G354370	细胞壁连接的类受体激酶5 Wall-associated receptor kinase 5-like

Genome-Wide Association Study of Powdery Mildew Resistance of Cucumber Core Germplasms

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