Association analysis of dynamic plant height trait using SSR marker in Gossypium hirsutum L.

doi:10.6048/j.issn.1001-4330.2023.12.005

Abstract

Abstract:

【Objective】To carry out association analysis of plant height of 185 upland cotton cultivars during the cotton flowering stage (PH-ST₁) and spikelike inflorescence period (PH-ST₂) to investigate the superior alleles variation of dynamic plant height. Which can provide a guidance for further high-yield molecular marker-assisted selection breeding of upland cotton.【Methods】355 polymorphism loci developed from 137 pairs of SSR polymorphism primers were combined with plant height phenotype data from two periods in five environments, and the general linear model (GLM) and mixed linear model (MLM) were used for association analysis by the genotypes of 185 upland cotton varieties.【Results】54 and 12 alleles in PH-ST₁ were detected by the GLM and MLM model, respectively. Meanwhile, 62 and 12 alleles in PH-ST₂ were detected by the GLM and MLM model. Among them, 31 loci were significantly related to plant height in 3 or more environments.【Conclusion】Through correlation analysis, multiple molecular markers that can be repeatedly detected and associated with the height of upland cotton were excavated.The 24 loci were reported.

Key words: Gossypium hirsutum L.; SSR; plant height; GLM; MLM

摘要：

【目的】分析185份陆地棉品种(系)的开花期株高(PH-ST₁)和吐絮期株高(PH-ST₂)关联度,发掘动态株高的优异等位变异,为进一步棉花高产分子辅助育种提供参考。【方法】利用185份陆地棉品种(系)的基因型,将137对简单重复序列(simple sequence repeat,SSR)多态性引物开发出的355个多态性位点,并结合5个环境下两个时期的株高表型数据,采用一般线性模型(general linear model,GLM)和混合线性模型(mixed linear model,MLM)进行关联分析。【结果】使用GLM模型和MLM模型,分别检测到54、12个与PH-ST₁显著相关的位点和62、12个与PH-ST₂显著相关的位点;其中,31个位点在3个或3个以上的环境都与株高显著相关。【结论】挖掘了多个可重复检测到的与陆地棉株高相关联的分子标记位点。定位到与株高性状相关的位点有24个。

关键词: 陆地棉, SSR, 株高, GLM, MLM

CLC Number:

S562
S188

WEN Jia, HUANG Chenjue, JI Zihan, LI Libei, FENG Zhen, YU Shuxun. Association analysis of dynamic plant height trait using SSR marker in Gossypium hirsutum L.[J]. Xinjiang Agricultural Sciences, 2023, 60(12): 2892-2901.

文佳, 黄陈珏, 嵇子涵, 李黎贝, 冯震, 喻树迅. 陆地棉动态株高与SSR标记的关联分析[J]. 新疆农业科学, 2023, 60(12): 2892-2901.

Figures/Tables 10

Tab.1 Statistical analysis of plant height traits in 185 upland cotton cultivars under different environments

环境 Environment	时期 Stage	均值 Mean	极小值 Min	极大值 Max	标准差 SD	变异系数 CV(%)	G	G×E	遗传率 h²(%)
中国海南三亚(2020年) Sanya,Hainan,China(2020)	开花期	58.45	35.86	80.84	8.72	14.56	^***	^***	63.67
中国湖北黄冈(2021年) Huanggang,Hubei China(2021)	开花期	46.93	34.41	59.63	5.84	12.45
中国海南三亚(2021年) Sanya,Hainan,China(2021)	开花期	60.02	36.67	84.47	7.95	13.29
中国山东临清(2021年) Linqing,Shandong,China(2021)	开花期	59.92	38.94	76.48	7.49	12.82
中国海南三亚(2020年) Sanya,Hainan,China(2020)	吐絮期	66.21	40.07	98.73	11.14	16.83	^***	^***	85.66
中国湖北黄冈(2021年) Huanggang,Hubei China(2021)	吐絮期	98.05	56.15	136.85	18.07	18.43
中国海南三亚(2021年) Sanya,Hainan,China(2021)	吐絮期	71.66	43.50	106.78	10.61	14.81
中国山东临清(2021年) Linqing,Shandong,China(2021)	吐絮期	88.30	53.15	123.41	15.78	17.87

Fig.1 The population structure of 185 upland cotton accessions

Tab.2 Association analysis results of PH-ST1 based on GLM model in 3 or more environments

位点 Locus	GLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
NAU828-418bp					2.35	4.16	3.14	5.56	2.67	4.41
DPL0524-3193bp	2.92	5.12			2.56	4.77	2.69	4.75	3.10	5.37
CGR5202-175bp	7.20	14.19			6.00	12.45			4.61	8.64
HAU2873-340bp	3.75	6.74			4.54	9.02	4.08	7.66	5.25	9.59
MGHES-73-352bp					2.34	4.26	5.68	10.92	3.60	6.36
HAU3318-310bp	4.41	7.98			5.24	10.51	4.96	9.43	5.97	10.95
HAU3318-308bp	5.07	9.30			4.81	9.60	5.92	11.36	6.69	12.33
PGML01548-281bp	3.60	6.48			3.15	6.00			4.15	7.50
NAU797-307bp	2.82	5.01			4.42	9.11			3.92	7.29
BNL2646-161bp	3.14	5.61	3.46	6.27	3.87	7.69	2.65	4.74	5.01	9.25
SWU0146-181bp			2.49	4.25	4.95	9.79	2.81	4.99	4.66	8.31
SWU0376-137bp			2.45	4.14	3.93	7.60			3.41	5.92
SWU0483-186bp					3.91	7.43	2.63	4.53	3.89	6.76
SWU0951-261bp	2.41	4.37	3.21	6.23	3.03	6.23			4.06	7.77
SWU1132-192bp	3.32	5.89			4.91	9.89			4.05	7.26
NAU2238-3			5.05	9.45	4.16	8.16	5.67	10.83	6.27	11.56
PGML01548-405bp	3.73	6.66	2.45	4.16					3.00	5.15
SWU0218-141bp			2.46	4.17			3.83	7.03	3.44	5.96
NAU4042-185bp	2.61	4.50					4.23	8.00	3.18	5.58

Tab.2 Association analysis results of PH-ST1 based on GLM model in 3 or more environments

位点 Locus	GLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
NAU828-418bp					2.35	4.16	3.14	5.56	2.67	4.41
DPL0524-3193bp	2.92	5.12			2.56	4.77	2.69	4.75	3.10	5.37
CGR5202-175bp	7.20	14.19			6.00	12.45			4.61	8.64
HAU2873-340bp	3.75	6.74			4.54	9.02	4.08	7.66	5.25	9.59
MGHES-73-352bp					2.34	4.26	5.68	10.92	3.60	6.36
HAU3318-310bp	4.41	7.98			5.24	10.51	4.96	9.43	5.97	10.95
HAU3318-308bp	5.07	9.30			4.81	9.60	5.92	11.36	6.69	12.33
PGML01548-281bp	3.60	6.48			3.15	6.00			4.15	7.50
NAU797-307bp	2.82	5.01			4.42	9.11			3.92	7.29
BNL2646-161bp	3.14	5.61	3.46	6.27	3.87	7.69	2.65	4.74	5.01	9.25
SWU0146-181bp			2.49	4.25	4.95	9.79	2.81	4.99	4.66	8.31
SWU0376-137bp			2.45	4.14	3.93	7.60			3.41	5.92
SWU0483-186bp					3.91	7.43	2.63	4.53	3.89	6.76
SWU0951-261bp	2.41	4.37	3.21	6.23	3.03	6.23			4.06	7.77
SWU1132-192bp	3.32	5.89			4.91	9.89			4.05	7.26
NAU2238-3			5.05	9.45	4.16	8.16	5.67	10.83	6.27	11.56
PGML01548-405bp	3.73	6.66	2.45	4.16					3.00	5.15
SWU0218-141bp			2.46	4.17			3.83	7.03	3.44	5.96
NAU4042-185bp	2.61	4.50					4.23	8.00	3.18	5.58

Tab.3 Association analysis results of PH-ST2 based on GLM model in 3 or more environments

位点 Locus	GLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
PGML02558-395bp					3.75	6.03	3.27	5.29	3.25	5.13
NAU4042-185bp	2.68	4.59			5.87	9.95	5.88	10.09	5.17	8.63
NAU3913-415bp					3.14	5.04	2.73	4.34	2.80	4.38
NAU1042-351bp					3.12	4.90	2.43	3.70	2.91	4.47
NAU833-433bp					3.86	6.12	2.70	4.13	3.44	5.33
NAU828-418bp					4.45	7.15	3.21	5.04	3.93	6.19
DPL0524-199bp					3.24	5.21	3.39	5.57	2.66	4.12
DPL0524-3193bp	3.18	5.58			4.82	8.07	4.37	7.36	4.70	7.77
BNL1705-180bp					2.57	3.87	2.68	4.16	2.79	4.26
HAU2873-340bp	3.99	7.13	2.38	4.11	7.09	12.02	6.20	10.59	6.90	11.55
MGHES-73-352bp					4.12	6.77	5.35	9.07	4.45	7.28
MGHES-73-340bp	2.39	4.11			4.26	7.28	2.41	3.85	2.96	4.79
NAU1102-236bp					3.25	5.22	4.70	7.92	4.00	6.51
NAU1190-222bp					2.39	3.58	3.96	6.42	3.04	4.69
HAU3318-310bp	3.52	6.19			7.53	12.78	6.68	11.44	6.12	10.23
HAU3318-308bp	4.40	7.92			6.96	11.81	7.33	12.57	6.64	11.11
PGML01548-281bp	3.15	5.50			3.73	6.10	3.25	5.28	3.72	6.03
NAU797-307bp	2.98	5.30			3.38	5.66	3.30	5.61	3.47	5.79
BNL2646-161bp	3.15	5.57	3.43	6.31	6.01	10.30	4.36	7.40	6.08	10.29
SWU0146-195bp					4.81	7.91	3.37	5.43	3.48	5.50
SWU0146-181bp			2.71	4.75	4.51	7.38	3.65	5.90	4.62	7.46
SWU0163-191bp					4.52	7.42	2.95	4.67	3.72	5.91
SWU0140-175bp			2.45	4.20	5.90	9.79	7.91	13.35	6.93	11.45
SWU0483-373bp			2.36	4.03	2.71	4.16	3.50	5.64	3.76	5.97
SWU0483-186bp			2.43	4.10	3.09	4.76	2.82	4.33	3.64	5.69
SWU0951-261bp			3.14	6.10	3.49	5.95	2.33	3.74	3.86	6.51
SWU0987-206bp					3.60	5.80	3.70	6.07	4.01	6.45
NAU2679-2					4.02	6.94	3.64	6.27	3.13	5.18
NAU2238-3			5.63	10.75	8.68	14.63	6.60	11.26	8.45	14.17
NAU2173-3					3.16	4.97	2.60	3.99	2.35	3.50
NAU2132					4.47	7.44	4.06	6.78	3.64	5.90
NAU1269-3					4.52	7.33	4.20	6.85	3.79	6.01
NAU1255-2					4.14	6.71	3.80	6.17	3.51	5.56
NAU1230-2					4.37	7.17	4.35	7.21	3.72	5.96

Tab.3 Association analysis results of PH-ST2 based on GLM model in 3 or more environments

位点 Locus	GLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
PGML02558-395bp					3.75	6.03	3.27	5.29	3.25	5.13
NAU4042-185bp	2.68	4.59			5.87	9.95	5.88	10.09	5.17	8.63
NAU3913-415bp					3.14	5.04	2.73	4.34	2.80	4.38
NAU1042-351bp					3.12	4.90	2.43	3.70	2.91	4.47
NAU833-433bp					3.86	6.12	2.70	4.13	3.44	5.33
NAU828-418bp					4.45	7.15	3.21	5.04	3.93	6.19
DPL0524-199bp					3.24	5.21	3.39	5.57	2.66	4.12
DPL0524-3193bp	3.18	5.58			4.82	8.07	4.37	7.36	4.70	7.77
BNL1705-180bp					2.57	3.87	2.68	4.16	2.79	4.26
HAU2873-340bp	3.99	7.13	2.38	4.11	7.09	12.02	6.20	10.59	6.90	11.55
MGHES-73-352bp					4.12	6.77	5.35	9.07	4.45	7.28
MGHES-73-340bp	2.39	4.11			4.26	7.28	2.41	3.85	2.96	4.79
NAU1102-236bp					3.25	5.22	4.70	7.92	4.00	6.51
NAU1190-222bp					2.39	3.58	3.96	6.42	3.04	4.69
HAU3318-310bp	3.52	6.19			7.53	12.78	6.68	11.44	6.12	10.23
HAU3318-308bp	4.40	7.92			6.96	11.81	7.33	12.57	6.64	11.11
PGML01548-281bp	3.15	5.50			3.73	6.10	3.25	5.28	3.72	6.03
NAU797-307bp	2.98	5.30			3.38	5.66	3.30	5.61	3.47	5.79
BNL2646-161bp	3.15	5.57	3.43	6.31	6.01	10.30	4.36	7.40	6.08	10.29
SWU0146-195bp					4.81	7.91	3.37	5.43	3.48	5.50
SWU0146-181bp			2.71	4.75	4.51	7.38	3.65	5.90	4.62	7.46
SWU0163-191bp					4.52	7.42	2.95	4.67	3.72	5.91
SWU0140-175bp			2.45	4.20	5.90	9.79	7.91	13.35	6.93	11.45
SWU0483-373bp			2.36	4.03	2.71	4.16	3.50	5.64	3.76	5.97
SWU0483-186bp			2.43	4.10	3.09	4.76	2.82	4.33	3.64	5.69
SWU0951-261bp			3.14	6.10	3.49	5.95	2.33	3.74	3.86	6.51
SWU0987-206bp					3.60	5.80	3.70	6.07	4.01	6.45
NAU2679-2					4.02	6.94	3.64	6.27	3.13	5.18
NAU2238-3			5.63	10.75	8.68	14.63	6.60	11.26	8.45	14.17
NAU2173-3					3.16	4.97	2.60	3.99	2.35	3.50
NAU2132					4.47	7.44	4.06	6.78	3.64	5.90
NAU1269-3					4.52	7.33	4.20	6.85	3.79	6.01
NAU1255-2					4.14	6.71	3.80	6.17	3.51	5.56
NAU1230-2					4.37	7.17	4.35	7.21	3.72	5.96

Tab.4 Association analysis results of PH-ST1 based on MLM model in 3 or more environments

位点 Locus	MLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
CGR5202-175bp	4.43	10.21			3.51	8.55			3.53	8.10
SWU0483-338bp	NA	0	NA	0	NA	0	NA	0	NA	0

Tab.4 Association analysis results of PH-ST1 based on MLM model in 3 or more environments

位点 Locus	MLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
CGR5202-175bp	4.43	10.21			3.51	8.55			3.53	8.10
SWU0483-338bp	NA	0	NA	0	NA	0	NA	0	NA	0

Tab.5 Association analysis results of PH-ST2 based on MLM model in 3 or more environments

位点 Locus	MLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
NAU2503-1			3.25	7.85			2.40	4.88	2.54	5.43
SWU0483-338bp	NA	0	NA	0	NA	0			NA	0

Tab.5 Association analysis results of PH-ST2 based on MLM model in 3 or more environments

位点 Locus	MLM
	中国海南三亚 (2020年) Sanya,Hainan, China(2020)		中国海南三亚 (2021年) Sanya,Hainan, China(2021)		中国湖北黄冈 (2021年) Huanggang,Hubei China(2021)		中国山东临清 (2021年) Linqing,Shandong, China(2021)		BLUP值
	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)	lg( $\bar{P}$ )	R²(%)
NAU2503-1			3.25	7.85			2.40	4.88	2.54	5.43
SWU0483-338bp	NA	0	NA	0	NA	0			NA	0

Fig.2 Venn diagrams of SSR locus associated with plant height at different stages based on GLM and MLM model

Fig.3 Phenotypic effects analysis of CGR5202-175bp locus Note:Hap 1:Material without the CGR5202-175bp locus;Hap 2:Material containing the CGR5202-175bp locus

Fig.4 Phenotypic effects analysis of NAU2238-3 locus Note:Hap 1:Material without NAU2238-3 locus;Hap 2:Material containing NAU2238-3 locus

Fig.5 Comparison of the QTL for plant height in cotton between this study and previous reports Note:Blue represents QTL in previous studies,red represents SSR loci within 1 Mb of QTL mapped previously

References 33

[1]	喻树迅. 与时俱进选育高产优质多抗短季棉品种[J]. 作物杂志, 2008,(6):1-2.
	YU Shuxun. Breeding short season cotton varieties with high yield,high quality and multi - resistance[J]. Crops, 2008,(6):1-2.
[2]	喻树迅. 我国棉花生产现状与发展趋势[J]. 中国工程科学, 2013, 15(4):9-13.
	YU Shuxun. Present situation and development trend of Cotton production in China[J]. Strategic Study of CAE, 2013, 15(4):9-13.
[3]	喻树迅, 张雷, 冯文娟. 快乐植棉——中国棉花生产的发展方向[J]. 棉花学报, 2015, 27(3):283-290. DOI
	YU Shuxun, ZHANG Lei, FENG Wenjuan. Easy and enjoyable cotton cultivation:developments in China’s cotton production[J]. Cotton Science, 2015, 27(3):283-290.
[4]	Qi H, Wang N, Qiao W, et al. Construction of a high-density genetic map using genotyping by sequencing(GBS)for quantitative trait loci(QTL)analysis of three plant morphological traits in upland cotton(Gossypium hirsutum L.)[J]. Euphytica, 2017, 213:83. DOI
[5]	Wang B H, Wu Y T, Huang N T, et al. QTL mapping for plant architecture traits in upland cotton using RILs and SSR markers[J]. Acta Genetica Sinica, 2006, 33(2):161-170. DOI URL
[6]	王新坤, 潘兆娥, 孙君灵, 等. 陆地棉矮秆突变体株高和纤维品质的QTL定位及相关性研究[J]. 核农学报, 2011, 25(3):448-455. DOI
	WANG Xinkun, PAN Zhao'e, SUN Junling, et al. Correlation analysis and QTL mapping for plant height and fiber quality of dwarf mutant in upland cotton (Gossypium hirsutum L.)[J]. Journal of Nuclear Agriculture, 2011, 25(3):448-455.
[7]	何蕊, 石玉真, 张金凤, 等. 利用染色体片段代换系定位陆地棉株高QTL[J]. 作物学报, 2014, 40(3):457-465.
	HE Rui, SHI Yuzhen, ZHANG Jinfeng, et al. QTL mapping for plant height using chromosome segment substitution lines in upland cotton[J]. Acta Agronomica Sinica, 2014, 40(3):457-465. DOI URL
[8]	Shang L, Fang L, Wang Y M, et al. Dynamic QTL mapping for plant height in Upland cotton (Gossypium hirsutum L.)[J]. Plant Breed, 2015, 134:703-712. DOI URL
[9]	Ma L L, Babar I, Wang Y M, et al. Dynamic qtl analysis and validation for plant height using maternal and paternal backcrossing populations in upland cotton[J]. Euphytica, 2018, 214(9):1-17. DOI
[10]	Nordborg M, Weigel D. Next-generation genetics in plants[J]. Nature, 2009, 456(7223):720-723. DOI
[11]	Gilad Y, Pritchard J K, Thornton K. Characterizing natural variation using next-generation sequencing technologies[J]. Trends in Genetics, 2009, 25(10):463-471. DOI PMID
[12]	Su J, Li L, Zhang C, Wang C, et al. Genome-wide association study identified genetic variations and candidate genes for plant architecture component traits in Chinese upland cotton[J]. Theoretical and Applied Genetics, 2018, 131(6):1299-1314. DOI PMID
[13]	Ji G, Liang C, Cai Y, et al. A copy number variant at the HPDA-D12 locus confers compact plant architecture in cotton[J]. The New Phytologist, 2021, 229(4):2091-2103. DOI URL
[14]	王晓东, 张洁夫, 栗茂腾. 甘蓝型油菜不同发育时期株高的动态QTL定位与比较分析[C]//. 江苏省遗传学会2014年学术研讨会会议论文摘要集.
	WANG Xiaodong, ZHANG Jiefu, LI Maoteng. Dynamic QTL mapping and comparative analysis of Plant height in Brassica napus[C]//. Abstract Collection of 2014 Academic Conference Papers of Jiangsu Genetics Society.
[15]	Shang L, Ma L, Wang Y, et al. Main Effect QTL with Dominance Determines Heterosis for Dynamic Plant Height in Upland Cotton[J]. G3(Bethesda), 2016, 6(10):3373-3379.
[16]	宋国立, 崔荣霞, 王坤波, 等. 改良CTAB法快速提取棉花DNA[J]. 棉花学报, 1998,(5):50-52.
	SONG Guoli, CUI Rongxia, WANG Kunbo, et al. A rapid improve CTAB method for extraction of cotton genomic DNA[J]. Cotton Science, 1998,(5):50-52.
[17]	刘其宝, 李黎贝, 张驰, 等. 陆地棉叶片叶绿素含量与SSR标记的关联分析及优异等位变异的挖掘[J]. 中国农业科学, 2017, 50(18):3439-3449. DOI
	LIU Qibao, LI Libei, ZHANG Chi, et al. Association analysis of leaf chlorophyll content with SSR markers and exploration of superior alleles in upland cotton[J]. Scientia Agricultura Sinica, 2017, 50(18):3439-3449. DOI
[18]	Breseghello F, Sorrells ME. Association mapping of kernel size and milling quality in wheat (Triticum aestivum L.) cultivars[J]. Genetics, 2006, 172(2):1165-1177. DOI PMID
[19]	Evanno G, Regnaut S, Goudet J. Detecting the number of clusters of individuals using the software STRUCTURE: a simulation study[J]. Molecular Ecology, 2005, 14(8):2611-2620. DOI PMID
[20]	Knapp S J, Stroup W W, Ross W M. Exact confidence intervals for heritability on a progeny mean basis[J]. Crop Science, 1985, 25:192-194. DOI URL
[21]	Bates D, Möchler M, Bolker B, et al. Fitting Linear Mixed-Effects Models Using lme4[J]. Journal of Statistical Software, 2015, 67(1):1-48.
[22]	Zhang B, Shi W, Li W, et al. Efficacy of pyramiding elite alleles for dynamic development of plant height in common wheat[J]. Molecular breeding, 2013, 32(2):327-338. DOI URL
[23]	Liu S, Saha S, Stelly D, et al. Chromosomal assignment of microsatellite loci in cotton[J]. The Journal of Heredity, 2000, 91(4):326-332. DOI URL
[24]	Gupta P, Varshney R. The development and use of microsatellite markers for genetic analysis and plant breeding with emphasis on bread wheat[J]. Euphytica, 2000, 113:163-185. DOI URL
[25]	Powell W, Machray G C, Provan J. Polymorphism revealed by simple sequence repeats[J]. Trends in Plant Science, 1996, 1(7):215-222. DOI URL
[26]	Li L, Zhang C, Huang J, et al. Genomic analyses reveal the genetic basis of early maturity and identification of loci and candidate genes in upland cotton(Gossypium hirsutum L.)[J]. Plant Biotechnology Journal, 2021, 19(1):109-123. DOI URL
[27]	Morris G P, Ramu P, Deshpande S P, et al. Population genomic and genome-wide association studies of agroclimatic traits in sorghum[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(2):453-458. DOI PMID
[28]	Su J, Pang C, Wei H, et al. Identification of favorable SNP alleles and candidate genes for traits related to early maturity via GWAS in upland cotton[J]. BMC Genomics, 2016, 17(1):687. DOI URL
[29]	Sun C, Zhang F, Yan X, et al. Genome-wide association study for 13 agronomic traits reveals distribution of superior alleles in bread wheat from the Yellow and Huai Valley of China[J]. Plant Biotechnology Journal, 2017, 15(8):953-969. DOI PMID
[30]	Wang J, Hu B, Jing Y, et al. Detecting QTL and Candidate Genes for Plant Height in Soybean via Linkage Analysis and GWAS[J]. Frontiers in Plant Science, 2022, 12:803820. DOI URL
[31]	Zhang Z, Ersoz E, Lai C Q, et al. Mixed linear model approach adapted for genome-wide association studies[J]. Nature Genetics, 2010, 42(4):355-360. DOI PMID
[32]	石桃雄, 唐彬, 任蓉蓉, 等. 苦荞AGPase编码基因FtAGPL和FtAGPS全基因组鉴定和表达分析[J]. 贵州师范大学学报(自然科学版), 2021, 39(4):52-57.
	SHI Taoxiong, TANG Bin, REN Rongrong, et al. Genome-wide identification and gene expression analysis of AGPase encoding genes FtAGPL and FtAGPS INTartary buckwheat (Fagopyrum tataricum)[J]. Journal of Guizhou Normal University (Natural Science Ed.), 2021, 39(4):52-57.
[33]	王龙, 李黎贝, 马启峰, 等. 陆地棉抗黄萎病性状与SSR标记的关联分析[J]. 新疆农业科学, 2018, 55(9):1569-1578. DOI
	WANG Long, LI Libei, MA Qifeng, et al. Association analysis of verticillum wilt resistance trait using SSR marker in Gossypium hirsutum L.[J]. Xinjiang Agricultural Sciences, 2018, 55(9):1569-1578. DOI