茶树新梢中香叶醇樱草糖苷含量的全基因组关联分析

doi:10.3724/SP.J.1006.2023.24173

Abstract

Abstract:

Geraniol is an important volatile monoterpene alcohol contained in tea plant, which plays an important role in the interaction between tea plant and environment and constitutes one of key aroma components in tea. Geraniol from tea plant tender shoots mainly exists in the form of geranyl β-primeveroside. Explore SNP loci and candidate genes significantly associated with geranyl β-primeveroside content is of great significance for the the genetic regulation mechanism of geranyl β-primeveroside content and tea plant genetic improvement. Regarding 169 tea germplasms as the research materials, the geranyl β-primeveroside content of tea plant tender shoots was identified for three consecutive years (three environments), and the SNP markers were developed based on SLAF-seq technology. Then, the geranyl β-primeveroside content of tea plant tender shoots was analyzed by genome-wide association analysis (GWAS) using general linear model (GLM). The candidate genes were further screened based on the SNP loci significantly with the geranyl β-primeveroside content. Ultimately, the base differences of candidate gene coding regions and their upstream cis acting elements among the extreme geranyl β-primeveroside content were analyzed. Results showed that the variation coefficient of geranyl β-primeveroside content in three environments ranged from 77.6% to 81.8%, and the broad heritability was 62.6%. The geranyl β-primeveroside content was significant differences among the genotypes and environments, and the variation was mainly affected by heredity. A total of 340 SNP markers significantly correlated with the geranyl β-primeveroside content were detected under three environments, in which 65 SNP markers were detected repeatedly under 2 environments. Based on the tea plant reference genome and linkage disequilibrium decay distance, a total of 88 genes within 100 kb on both sides of the repeated 65 SNP markers were obtained, including signal proteins, kinases, phosphatases, ion transporters, transcription factors, heat shock proteins, hormone related proteins, resistance proteins, terpene metabolic enzymes, glycosyltransferases, and glycosidases. 10 candidate genes were preliminary screened out. SNP nonsynonymous mutations in the coding region sequences of 10 candidate genes between two extreme geranyl β-primeveroside content materials, and different numbers of cis acting elements related to environmental stress and hormones in the upstream 2 kb regions of most candidate genes were discovered. This study provides a new perspective for clarifying the genetic regulation mechanism of geranyl β-primeveroside content in tea plant tender shoots, and also provides the markers and gene resources for molecular marker assisted selection of new tea varieties.

Key words: tea (Camellia sinensis L.), geranyl β-primeveroside, genome-wide association analysis (GWAS), single nucleotide polymorphism (SNP), candidate genes

WANG Rang-Jian, YANG Jun, ZHANG Li-Lan, GAO Xiang-Feng. Genome-wide association analysis of geraniol primrose glycoside abundance in tender tea shoots[J].Acta Agronomica Sinica, 2023, 49(7): 1843-1859.

Figures/Tables 13

Table 1

Table 2

Table 3

Fig. 1

Table 4

SNP sites related to geranyl β-primeveroside content"

染色体 Scaffold	SNP位置 SNP location	SNP变异 SNP variation	功能区 Functional area	贡献率 Contribution rate (%)	P值P-value
染色体 Scaffold	SNP位置 SNP location	SNP变异 SNP variation	功能区 Functional area	贡献率 Contribution rate (%)	2018	2019	2020
Scaffold106	940,480	C/T	间区Intergenic	19.3	—	4.25E-09	7.89E-08
	940,538	A/G	间区Intergenic	18.1	—	1.01E-08	1.41E-07
Scaffold1262	982,460	G/A	间区Intergenic	18.2	—	5.56E-09	6.45E-08
	982,581	A/T	间区Intergenic	18.2	—	5.56E-09	6.45E-08
Scaffold1318	1,188,898	C/A	间区Intergenic	20.7	2.70E-08	9.11E-10	—
Scaffold1611	610,786	C/T	间区Intergenic	20.7	1.60E-07	9.52E-09	—
Scaffold1671	583,382	C/T	内含子Intron	18.4	—	4.92E-09	7.52E-08
	583,566	G/A	内含子Intron	18.2	—	6.24E-09	8.42E-08
Scaffold209	336,614	G/A	间区Intergenic	22.6	—	1.88E-09	6.50E-08
Scaffold211	1,175,079	G/A	间区Intergenic	19.2	1.56E-07	8.21E-09	—
	1,175,109	T/A	间区Intergenic	19.2	1.56E-07	8.21E-09	—
	1,175,111	C/T	间区Intergenic	19.2	1.56E-07	8.21E-09	—
	1,175,135	C/T	间区Intergenic	19.2	1.56E-07	8.21E-09	—
	1,175,177	C/T	间区Intergenic	19.2	1.55E-07	8.32E-09	—
Scaffold2268	2,777,884	G/A	间区Intergenic	19.4	1.33E-07	1.43E-08	—
	2,777,990	G/A	间区Intergenic	20.4	1.26E-07	4.63E-09	—
Scaffold2311	361,209	C/T	间区Intergenic	18.5	1.00E-07	3.44E-09	—
Scaffold2800	1,171,675	A/T	间区Intergenic	19.8	8.69E-08	6.93E-09	—
Scaffold2942	113,797	G/A	间区Intergenic	20.0	1.15E-07	1.00E-08	—
	113,806	C/T	间区Intergenic	20.0	1.15E-07	1.00E-08	—
Scaffold30	1,872,412	G/T	间区Intergenic	20.2	—	6.24E-09	7.68E-08
Scaffold3229	195,605	C/A	间区Intergenic	17.1	—	1.56E-08	1.53E-07
	1,193,442	C/T	间区Intergenic	20.7	—	2.28E-09	3.86E-09
Scaffold3317	78,232	C/A	内含子Intron	23.9	—	2.35E-10	7.93E-10
	78,432	G/A	内含子Intron	21.8	—	2.12E-09	1.47E-09
Scaffold3395	141,462	C/T	间区Intergenic	20.4	8.41E-09	3.16E-10
	141,471	G/A	间区Intergenic	20.4	8.41E-09	3.16E-10
	141,479	G/A	间区Intergenic	20.4	8.41E-09	3.16E-10
	141,537	G/A	间区Intergenic	20.4	8.41E-09	3.16E-10
Scaffold349	3,542,803	G/A	间区Intergenic	21.5	8.21E-09	4.86E-10
	3,543,009	G/A	间区Intergenic	22.0	4.03E-09	3.10E-10
	3,543,014	G/A	间区Intergenic	21.0	6.06E-09	9.46E-10
	3,543,021	G/T	间区Intergenic	20.8	8.62E-09	1.16E-09
	3,543,061	C/T	间区Intergenic	21.0	7.19E-09	9.75E-10
	3,543,082	G/A	间区Intergenic	21.0	6.52E-09	9.75E-10
Scaffold3611	1,622,374	A/C	间区Intergenic	18.7	—	9.80E-09	8.04E-08
	1,622,376	A/G	间区Intergenic	18.7	—	9.80E-09	8.04E-08
Scaffold4122	389,378	G/A	间区Intergenic	19.5	—	2.34E-09	9.22E-10
	444,511	C/T	间区Intergenic	18.8	—	2.51E-09	7.63E-10
	444,749	C/A	间区Intergenic	18.8	—	2.51E-09	7.63E-10
	444,760	G/A	间区Intergenic	19.4	—	2.37E-09	7.33E-10
Scaffold4659	1,061,780	G/A	间区Intergenic	19.0	3.62E-08	2.72E-09	—
Scaffold4817	276,251	C/T	间区Intergenic	17.8	6.68E-08	5.92E-09	—
Scaffold522	69,932	T/C	内含子Intron	20.2	8.67E-08	6.37E-09	—
	69,936	T/C	内含子Intron	20.2	8.67E-08	6.37E-09	—
Scaffold708	2,098,580	G/A	间区Intergenic	19.8	—	7.24E-09	1.21E-07
	2,098,607	G/A	间区Intergenic	20.4	—	4.00E-09	6.13E-08
	2,098,614	C/T	间区Intergenic	19.9	—	7.01E-09	1.20E-07
	2,098,617	G/A	间区Intergenic	19.8	—	7.24E-09	1.21E-07
	2,098,661	A/G	间区Intergenic	19.5	—	8.39E-10	1.67E-08
	2,098,666	G/C	间区Intergenic	18.0	—	4.86E-09	5.84E-08
	2,098,674	G/A	间区Intergenic	18.0	—	4.86E-09	5.84E-08
	2,098,695	T/C	间区Intergenic	18.0	—	4.86E-09	5.84E-08
Scaffold7165	292,911	A/T	下游Downstream	22.7	9.53E-09	7.52E-10	—
Scaffold8160	383,511	C/T	内含子Intron	18.6	8.16E-08	5.10E-09	—
Scaffold8614	526,707	A/G	间区Intergenic	20.8	7.33E-08	3.20E-09	—
	526,802	C/T	间区Intergenic	19.0	—	1.36E-09	5.85-08
Scaffold8631	103,446	G/A	间区Intergenic	21.9	—	1.29E-09	2.71E-08
	103,448	T/C	间区Intergenic	21.7	—	1.73E-09	3.43E-08
	103,743	G/A	间区Intergenic	21.9	—	1.29E-09	2.71E-08
Scaffold877	1,740,414	C/T	间区Intergenic	19.8	—	1.01E-08	1.39E-07
	1,740,416	G/A	间区Intergenic	22.0	—	1.00E-09	2.50E-08
	1,740,418	G/A	间区Intergenic	21.2	—	2.37E-09	2.12E-08
Scaffold89	2,029,626	A/G	内含子 Intron	19.1	—	7.64E-09	1.18E-07
Scaffold9179	390,469	A/T	内含子 Intron	19.4	—	9.31E-09	1.22E-07

Table 4

Table 5

Candidate gene information"

染色体 Scaffold	SNP位置 SNP location	距离 Distance (kb)	候选基因 Candidate gene	功能注释 Function annotation
Scaffold106	940,480-940,538	5°-37.4	TEA029079.1	MYB转录因子MYB Transcription factor
		5°-65.0	TEA029081.1	丝氨酸/苏氨酸蛋白磷酸酶Serine/threonine-protein kinase
		5°-76.2	TEA029085.1	脱落酸受体PYL2 Abscisic acid receptor PYL2
Scaffold1262	982,460-982,581	3°-29.7	TEA016845.1	MYB转录因子MYB Transcription factor
Scaffold1318	1,188,898	5°-73.4	TEA023905.1	PTB 调控因子PTB Regulation factor
		5°-58.4	TEA023908.1	亮氨酸重复受体蛋白激酶 Leucine-rich repeat receptor-like protein kinase
		5°-68.5	TEA023909.1	亮氨酸重复受体蛋白激酶 Leucine-rich repeat receptor-like protein kinase
		5°-11.7	TEA023910.1	莽草酸邻羟基肉桂酰转移酶 Shikimate O-hydroxy cinnamoyl transferase
Scaffold1611	610,716-610,786	5°-17.0	TEA001327.1	丝氨酸/苏氨酸蛋白激酶Serine/threonine-protein kinase
Scaffold1671	583,382-583,566	5°-47.6	TEA026622.1	抗病蛋白Disease resistance protein
		3°-87.8	TEA026619.1	乙烯反应转录因子 Ethylene-responsive transcription factor ERF
		5°-53.1	TEA026637.1	抗病蛋白Disease resistance protein
Scaffold209	336,614	5°-99.2	TEA015065.1	半乳糖醛酸转移酶Galacturonosyltransferase
		5°-44.9	TEA015072.1	丝氨酸/苏氨酸蛋白激酶Serine/threonine-protein kinase
Scaffold211	1,175,079-1,175,177	3°-66.4	TEA033378.1	糖基转移酶Glycosyltransferase
		5°-37.2	TEA033379.1	细胞色素P450 Cytochrome P450
		3°-12.1	TEA033380.1	未知蛋白Uncharacterized protein
		3°-21.8	TEA033381.1	MOR1 蛋白MOR1 Protein
		3°-6.6	TEA033375.1	未知蛋白Uncharacterized protein
Scaffold2268	2,777,884-2,777,990	5°-53.5	TEA021079.1	抗病蛋白Disease resistance protein
		5°-59.0	TEA021105.1	亮氨酸重复受体蛋白激酶 Leucine-rich repeat receptor-like protein kinase
		5°-95.5	TEA021068.1	TMV抗性蛋白TMV Resistance protein
Scaffold2311	361,209	5°-79.8	TEA030505.1	丝氨酸/苏氨酸蛋白激酶Serine/threonine-protein kinase
		5°-42.6	TEA030506.1	未知蛋白Uncharacterized protein
		5°-7.7	TEA030509.1	未知蛋白Uncharacterized protein
Scaffold2800	1,171,675	3°-23.5	TEA002738.1	钙转运ATP酶Calcium-transporting ATPase
		5°-1.6	TEA002761.1	信号蛋白Signalling protein
		5°-7.7	TEA002758.1	信号蛋白Signalling protein
		5°-67.9	TEA002766.1	抗性蛋白Resistance protein
		5°-19.5	TEA002755.1	丝氨酸羧肽酶Serine carboxypeptidase
Scaffold2942	113,797-113,806	5°-94.2	TEA018613.1	F-盒子调控蛋白F-box protein
Scaffold30	1,872,412	5°-32.2	TEA009798.1	淀粉合成酶Starch synthase
		5°-71.1	TEA009811.1	淀粉合成酶Starch synthase
		3°-74.6	TEA009793.1	叶绿素结合蛋白Chlorophyll a-b binding protein
Scaffold3229	1,195,605	3°-15.3	TEA007727.1	微管结合蛋白Microtubule binding protein
		3°-84.1	TEA007735.1	微管结合蛋白Microtubule binding protein
		5°-36.8	TEA007726.1	dof 转录因子dof transcription factor
Scaffold3317	78,232-78,432	5°-18.7	TEA014251.1	赤霉素氧化酶Gibberellin oxidase
		5°-43.8	TEA014250.1	膜相关激酶调节器Membrane-associated kinase regulator
Scaffold3395	141,462-141,537	5°-29.2	TEA023356.1	未知蛋白Uncharacterized protein
		3°-44.7	TEA023352.1	热激蛋白Heat shock 70 kD protein
Scaffold349	3,542,803-3,543,082	5°-8.6	TEA023826.1	异戊烯基二磷酸合酶/异戊烯基转移酶 Isoprenyl diphosphate synthase/prenyltransferase
		5°-10.9	TEA023811.1	核糖体蛋白Ribosomal protein
		5°-33.9	TEA023806.1	翻译起始因子Translation initiation factor
		5°-85.8	TEA023801.1	RNA聚合酶II转录亚基 RNA Polymerase II transcription subunit
		3°-89.3	TEA023809.1	RNA聚合酶II转录亚基 RNA Polymerase II transcription subunit
Scaffold3611	1,622,374-1,622,376	5°-4.5	TEA013393.1	WRKY转录因子WRKY Transcription factor
		3°-14.9	TEA013410.1	UDP-糖基转移酶 92A1 UDP-glycosyltransferase 92A1
Scaffold4122	444,511-444,760	3°-56.3	TEA027152.1	F-盒子调控蛋白F-box protein
		3°-69.0	TEA027153.1	F-盒子调控蛋白F-box protein
Scaffold4659	1,061,780	3°-30.6	TEA001887.1	细胞壁相关受体激酶Wall-associated receptor kinase
		5°-87.4	TEA001888.1	Beta-1,3-葡聚糖酶Beta-1,3-glucanase
Scaffold4817	276,251	3°-71.7	TEA006075.1	富含亮氨酸重复序列的蛋白激酶 Leucine-rich receptor-like protein kinase
		5°-86.7	TEA006076.1	抗病蛋白RPM1 Disease resistance protein RPM1
		5°-30.9	TEA006078.1	线粒体蛋白Mitochondria protein
Scaffold522	69,932-69,936	3°-11.3	TEA004895.1	抗病蛋白Disease resistance protein
		3°-50.6	TEA004897.1	转录因子ORG2 Transcription factor ORG2
		3°-12.1	TEA004887.1	未知蛋白Uncharacterized protein
Scaffold708	2,098,580-2,098,695	3°-40.7	TEA027473.1	MLO蛋白MLO-like protein
		5°-19.6	TEA027474.1	果胶酯酶/果胶酯酶抑制剂 Pectinesterase/pectinesterase inhibitor
		3°-92.8	TEA027481.1	果胶酯酶抑制剂Pectinesterase inhibitor
		3°-58.4	TEA027489.1	未知蛋白Uncharacterized protein
		5°-54.2	TEA027491.1	肽基脯氨酰顺反异构酶 Peptidylprolyl cis/trans isomerase
		5°-65.8	TEA027494.1	uncharacterized protein
		3°-74.6	TEA027468.1	转录激活因子Transcriptional activator
		5°-66.7	TEA027497.1	未知蛋白Uncharacterized protein
Scaffold7165	292,911	3°-4.3	TEA009868.1	钙转运ATP酶Calcium-transporting ATPase
		5°-4.5	TEA009869.1	ADP核糖基化因子ADP-ribosylation factor
Scaffold8160	383,511	3°-21.1	TEA009860.1	富含脯氨酸受体样蛋白激酶 Proline-rich receptor-like protein kinase
		3°-89.4	TEA009862.1	阳离子逆向转运蛋白Cation/H(+) antiporter
		5°-26.1	TEA009861.1	抗病蛋白Disease resistance protein
		3°-15.9	TEA009863.1	未知蛋白Uncharacterized protein
		3°-12.9	TEA009864.1	核糖体蛋白Ribosomal protein
		3°-14.1	TEA009865.1	亚硫酸盐输出蛋白Sulfite exporter protein
Scaffold8614	526,707-526,802	5°-49.9	TEA004103.1	钙调素结合蛋白Calmodulin-binding protein
Scaffold8631	103,446-103,743	3°-8.8	TEA006901.1	Ras相关蛋白Ras-related protein
		5°-8.5	TEA006902.1	Ras相关蛋白Ras-related protein
		3°-70.9	TEA006898.1	未知蛋白Uncharacterized protein
Scaffold877	1,740,414-1,740,418	3°-12.5	TEA030188.1	铜离子转运蛋白Copper transporter
		3°-46.8	TEA030196.1	阳离子/钙交换剂Cation/calcium exchanger
		5°-45.1	TEA030182.1	鸟嘌呤核苷酸结合蛋白Guanine nucleotide-binding protein
		3°-61.6	TEA030186.1	枯草杆菌蛋白酶Subtilisin-like protease
		3°-47.5	TEA030202.1	阳离子/钙交换剂Cation/calcium exchanger
		5°-8.7	TEA030204.1	未知蛋白Uncharacterized protein
		3°-86.2	TEA030207.1	蛋白酶体亚基Proteasome subunit
Scaffold89	2,029,626	5°-86.0	TEA018925.1	Beta-1,3-葡聚糖酶Beta-1,3-glucanase
Scaffold9179	390,469	3°-36.5	TEA027247.1	丝氨酸/苏氨酸蛋白激酶Serine/threonine-protein kinase
		3°-89.2	TEA027245.1	醛酮还原酶Aldo-keto reductase

Table 5

Fig. 2

Fig. 3

Fig. 4

Table 6

Table 7

Table 8

Fig. 5

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环境 Environment	均值 Mean value	标准差 Standard deviation	变异系数 Coefficient of variation (%)	Ryan-Joiner检测 Ryan-Joiner test
2018	0.3585	0.2809	78.4	r=0.917, P<0.01
2019	0.2674	0.2187	81.8	r=0.914, P<0.01
2020	0.3581	0.2779	77.6	r=0.855, P<0.01

方差来源 Variance	自由度 DF	平方和 SS	均方 MS	F值 F-value	遗传力h² Heritability h²(%)
基因 Genotype	168	34.1299	0.2032	6.12^**	62.6
环境 Environment	2	0.9318	0.4659	14.04^**
误差 Error	336	11.3233	0.0337

主要类型 Main types	最大数 Maximum number	最小数 Minimum number	平均数 Average number
基因间区Intergenic	1,595,407	513,424	1,062,687
内含子Intron	53,116	21,509	41,091
基因上游(5 kb内) Upstream in 5 kb	46,686	17,538	37,335
基因下游(5 kb内) Downstream in 5 kb	42,110	16,067	32,681
同义突变Synonymous	5004	2161	3537
非同义突变Nonsynonymous	6703	2931	4911
其他Other	2899	1161	2030

染色体 Scaffold	候选基因 Candidate gene	基因注释 Gene annotation	每百万比对数目Counts per million
染色体 Scaffold	候选基因 Candidate gene	基因注释 Gene annotation	409	1605	悦茗香^* Yuemingxiang^*	福云6号^* Fuyun 6^*
Scaffold1318	TEA023908.1	富含亮氨酸重复类受体蛋白激酶 Leucine-rich repeat receptor-like protein kinase	7.09	2.31	16.88	1.22
Scaffold4817	TEA006075.1	富含亮氨酸重复类受体蛋白激酶 Leucine-rich repea treceptor-like protein kinase	2.73	0.23	3.89	0.31
Scaffold106	TEA029081.1	丝氨酸/苏氨酸蛋白激酶 Serine/threonine-protein kinase	20.21	8.70	17.13	6.32
Scaffold9179	TEA027247.1	丝氨酸/苏氨酸蛋白激酶 Serine/threonine-protein kinase	11.93	5.26	22.37	4.63
Scaffold349	TEA023811.1	核糖体蛋白 Ribosomal protein	12.40	4.89	21.10	5.69
Scaffold8160	TEA009864.1	核糖体蛋白 Ribosomal protein	65.70	24.41	109.79	33.77
Scaffold349	TEA023801.1	RNA聚合酶II转录亚基 RNA polymerase II transcription subunit	139.33	70.50	147.46	59.41
Scaffold4659	TEA001888.1	Beta-1,3-葡聚糖酶 Beta-1,3-glucanase	7.65	1.09	8.94	1.92
Scaffold89	TEA018925.1	Beta-1,3-葡聚糖酶 Beta-1,3-glucanase	236.17	130.12	190.25	150.34
Scaffold4817	TEA006076.1	抗病蛋白RPM1 Disease resistance protein RPM1	12.33	0.85	13.07	1.31

候选基因 Candidate gene	SNP位置 SNP location	409 DNA序列 409 DNA sequence	1605 DNA序列 1605 DNA sequence	409氨基酸序列 409 amino acid sequence	1605氨基酸序列 1605 amino acid sequence
TEA006075.1	348,400	T/A	T	Lys/Ter	Lys
	348,421	C/G	C	Glu/Gln	Glu
	348,436	G/C	G	Leu/Val	Leu
	348,467	G/A	G	Ala/Val	Ala
	348,488	A/G	A	Met/Thr	Met
	348,524	C/T	C	Gly/Glu	Gly
	348,764	C/T	C	Ser/Asn	Ser
	348,797	C/T	C	Cys/Tyr	Cys
	348,830	G/A	G	Ser/Phe	Ser
	349,130	G/A	G	Ser/Phe	Ser
	349,248	A/T	A	Tyr/Ter	Tyr
	349,283	G/A	G	Ser/Phe	Ser
	349,370	T/A	T	His/Leu	His
	349,421	T/C	T	Ter/Trp	Ter
	349,536	G/C	G	Phe/Leu	Phe
	349,550	A/T	A	Ile/Lys	Ile
	349,625	C/T	C	Trp/Ter	Trp
TEA006075.1	349,633	T/G	T	Asn/His	Asn
	350,501	C/T	C	Trp/Ter	Trp
	350,647	G/A	G	Thr/Ile	Thr
TEA023908.1	1,127,395	A	G	Ile	Thr
	1,127,410	G	G/A	Ala	Ala/Val
	1,127,793	C/G	C	Gln	Glu
	1,127,867	T/A	T	Ser	Arg
	1,127,989	C/T	C	Asn	Ser
	1,128,205	A/G	A	Ala	Val
	1,128,366	G	G/C	Gln	Gln/Glu
	1,128,407	G	G/T	Phe	Phe/Leu
	1,128,445	A	A/G	Ile	Ile/Thr
	1,128,890	T	T/C	Ile	Ile/Met
	1,129,004	G/T	T	His/Gln	Gln
	1,130,137	A	A/G	Ile	Ile/Thr
	1,130,398	A/G	A	Ile/Thr	Ile
TEA029081.1	867,063	A/G	A	Leu/Ser	Leu
	867,147	C/T	T	Ser/Asn	Asn
	867,194	G	C	Pro	Ala
	867,221	C	A	Val	Leu
	867,354	A/G	A	Leu/Ser	Leu
	869,008	C	G	Arg	Thr
	869,038	T/A	T	His/Leu	His
	874,334	T	G	Lys	Gln
TEA006076.1	187,415	C/A	C	Leu/Phe	Leu
	187,442	G/T	G	Cys/Ter	Cys
	189,274	C/G	C	Cys/Ser	Cys
	189,283	T/C	T	Gln/Arg	Gln
TEA023801.1	3,449,354	G/C	C	Tyr/Ter	Ter
TEA023801.1	3,453,357	A	A/G	Leu	Leu/Pro
TEA023811.1	3,531,058	T	T/A	Lys	Lys/Ter
TEA001888.1	1,154,774	T	T/G	Met	Met/Leu
TEA009864.1	370,433	G/A	A	Ala/Val	Val
TEA027247.1	358,366	A	A/T	Phe	Phe/Ile
TEA018925.1	1,943,138	C	A	Arg	Ser

Genome-wide association analysis of geraniol primrose glycoside abundance in tender tea shoots

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候选基因 Candidate gene	激素响应元件 Hormone responsive elements					环境胁迫相关元件 Environmental stress-related elements
候选基因 Candidate gene	IAA	GA	SA	ABA	MeJA	ASI	WR	LTR	DSR	AI	DI
TEA029081.1			2		2			1		3	2
TEA023908.1	1		2		2				1	2
TEA023801.1								1		1
TEA023811.1		2		4						1
TEA001888.1	1	1					1		1	4
TEA006075.1		2	1	5	4			1	1	5
TEA006076.1	1	1		1	4				1
TEA009864.1
TEA018925.1	1	2			2	1			2	1	1
TEA027247.1				3	4			1		3	2

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