Mapping and Analysis of QTLs for Rice Grain Weight and Grain Shape Using Chromosome Segment Substitution Line Population

doi:10.16819/j.1001-7216.2021.210204

Abstract

Abstract:

【Objective】 Mining quantitative trait loci (QTL) for grain weight and grain shape related traits in rice plays an important role in the genetic mechanism of rice grain.【Method】 QTLs associated with grain weight and grain shape of grain and brown rice were analyzed using the population of chromosome segment substitution lines (CSSLs) with indica rice 9311 as recipient and japonica Nipponbare as donor in four different environments.【Result】 A total of 77 QTLs for grain weight and grain shape of paddy and brown rice were detected, and the phenotypic variation explained by each QTL ranged from 4.62% to 51.01%. Among these QTLs, 19 have additive alleles from Nipponbare and 58 have additive alleles from 9311. These QTLs distributed in 46 regions of 10 chromosomes in rice, and 16 regions are pleiotropic loci. There are 14 QTLs that can be repeatedly detected in two or more environments, qGW5.1 and qLW5 could be repeatedly detected in four environments and were located in the same chromosome region; qBRL3.2 and qGL4.1 were newly identified QTLs affecting grain weight and grain shape in rice.【Conclusion】 The results laid a foundation for cloning these QTLs and analyzing the genetic mechanism of grain shape and grain weight in rice.

Key words: rice, chromosome segment substitution lines (CSSLs), grain weight, grain shape, QTL

摘要：

【目的】挖掘水稻粒重和粒型相关性状QTL,对于解析水稻籽粒遗传机理具有重要作用。【方法】本研究以籼稻9311为受体、粳稻日本晴为供体构建的染色体片段置换系（Chromosome Segment Substitution Lines, CSSLs）群体为材料,在4个环境下对控制稻谷与糙米的粒重和粒型QTL进行了定位分析。【结果】共检测到77个控制水稻粒重和粒型的QTL,贡献率为4.62%~51.01%,其中19个QTL的增效等位基因来自日本晴,58个QTL的增效等位基因来自9311。这些QTL分布在水稻10条染色体的46个区域,其中16个区域为多效性位点。在两个及两个以上环境中重复检测到的QTL有14个,其中qGW5.1和qLW5能够在4个环境中稳定表达,且位于同一染色体区域;qBRL3.2和qGL4.1为新鉴定的影响水稻粒重和粒型的QTL。【结论】本研究结果为后续克隆这些QTL和解析水稻粒重和粒型遗传机理奠定了基础。

关键词: 水稻, 染色体片段置换系, 粒重, 粒型, QTL

HUANG Tao, WANG Yanning, ZHONG Qi, CHENG Qin, YANG Mengmeng, WANG Peng, WU Guangliang, HUANG Shiying, LI Caijing, YU Jianfeng, HE Haohua, BIAN Jianmin. Mapping and Analysis of QTLs for Rice Grain Weight and Grain Shape Using Chromosome Segment Substitution Line Population[J]. Chinese Journal OF Rice Science, 2022, 36(2): 159-170.

黄涛, 王燕宁, 钟奇, 程琴, 杨朦朦, 王鹏, 吴光亮, 黄诗颖, 李才敬, 余剑峰, 贺浩华, 边建民. 利用染色体片段置换系群体定位和分析水稻粒重和粒型QTL[J]. 中国水稻科学, 2022, 36(2): 159-170.

Figures/Tables 7

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性状 Trait	环境Environment	亲本 Parent		置换系群体 CSSL
性状 Trait	环境Environment	9311	日本晴 Nipponbare	均值±标准Mean±SD	变幅 Range	峰度Kurtosis	偏度Skewness
稻谷粒长	E1	9.86±0.47**	7.38±1.56	9.59±0.35	8.37~10.69	-0.46	2.58
Grain length / mm	E2	9.76±0.08**	7.68±0.16	9.31±0.28	8.23~9.77	-1.66	3.99
	E3	9.79±0.16**	7.67±0.19	9.52±0.33	8.05~10.14	-1.79	5.19
	E4	9.80±0.24**	7.63±0.29	9.78±0.31	8.71~10.30	-1.06	1.42
稻谷粒宽	E1	2.79±0.12*	3.24±0.60	2.78±0.13	2.55~3.53	2.33	10.17
Grain width / mm	E2	2.48±0.12*	3.30±0.56	2.99±0.12	2.69~3.61	2.00	8.05
	E3	2.59±0.19*	3.17±0.00	2.81±0.13	2.43~3.38	1.83	6.11
	E4	2.65±0.14*	3.13±0.42	2.83±0.11	2.56~3.27	1.43	4.11
稻谷长宽比	E1	3.55±0.27**	2.29±0.13	3.49±0.19	2.62~3.78	-1.91	4.62
Length-width ratio	E2	3.47±0.12**	2.31±0.23	3.14±0.16	2.42~3.38	1.60	3.70
	E3	3.78±0.11**	2.42±0.19	3.41±0.19	2.71~3.71	-1.59	2.73
	E4	3.70±0.05**	2.34±0.14	3.47±0.18	2.60~3.80	-1.82	5.04
稻谷粒厚	E1	2.26±0.09	2.17±0.05	2.19±0.05	2.04~2.34	0.07	0.64
Grain thickness / mm	E2	2.14±0.10	2.09±0.06	2.18±0.05	2.05~2.33	0.20	0.12
	E3	2.09±0.05	2.19±0.09	2.12±0.06	1.93~2.24	-0.54	0.05
	E4	1.99±0.06	2.01±0.12	2.10±0.05	1.95~2.20	-0.33	0.05
稻谷千粒重	E1	31.02±2.37**	22.44±1.69	29.51±2.29	22.17~34.21	-0.64	0.70
1000-grain weight / g	E2	31.36±3.06**	22.26±2.01	29.63±1.97	23.24~34.90	-0.40	0.77
	E3	30.39±2.74**	21.86±1.49	27.39±2.42	19.71~33.65	-0.21	0.48
	E4	32.53±2.36**	22.65±1.84	29.46±2.16	23.46~33.49	-0.58	0.41
糙米长	E1	6.87±0.31**	4.88±0.36	6.91±0.23	6.14~7.30	-1.50	2.67
Brown rice length	E2	6.84±0.41**	0.86±0.49	6.84±0.24	5.94~7.36	-1.64	3.57
/mm	E3	6.91±0.46**	4.99±0.61	6.93±0.24	5.96~7.39	-1.54	3.65
	E4	6.91±0.37**	4.70±0.23	6.92±0.27	5.97~7.54	-0.57	0.55
糙米宽	E1	2.38±0.12*	2.70±0.08	2.36±0.11	2.17~2.92	2.04	7.85
Brown rice width	E2	2.27±0.14*	2.67±0.13	2.52±0.09	2.31~3.01	1.90	9.25
/mm	E3	2.32±0.19*	2.78±0.05	2.40±0.09	2.16~2.79	1.50	5.41
	E4	2.46±0.11*	2.75±0.06	2.36±0.11	2.07~2.89	1.17	4.67
糙米长宽比	E1	2.98±0.21**	1.87±0.19	2.95±0.17	2.17~3.21	-1.67	3.84
Length-to-width ratio	E2	2.94±0.23**	1.84±0.17	2.72±0.14	2.11~2.99	-1.56	4.07
of brown rice	E3	2.98±0.31**	1.81±0.16	2.91±0.15	2.33~3.15	-1.53	3.09
	E4	2.76±0.27**	1.86±0.19	2.95±0.14	2.26~3.16	-1.95	5.56
糙米厚	E1	1.93±0.10	2.07±0.06*	1.93±0.05	1.79~2.10	0.14	0.66
Thickness of brown rice	E2	1.90±0.01	2.12±0.04*	1.91±0.05	1.77~2.08	0.36	1.48
/ mm	E3	1.85±0.07	1.96±0.06*	1.87±0.05	1.73~2.07	0.10	2.63
	E4	1.91±0.06	2.10±0.05*	1.88±0.05	1.77~2.06	0.37	0.38
糙米千粒重	E1	23.55±0.49*	21.88±0.32	24.64±1.57	20.47~29.46	0.03	0.33
1000-grain weight	E2	22.86±0.09*	21.76±0.36	24.32±1.37	20.34~28.20	-0.21	0.72
of brown rice / g	E3	22.29±0.14*	20.35±0.60	23.14±1.61	18.19~27.60	-0.10	0.31
	E4	23.67±0.52*	21.66±0.54	2.83±1.56	19.63~26.95	-0.39	0.00

性状 Trait	环境Environment	亲本 Parent		置换系群体 CSSL
性状 Trait	环境Environment	9311	日本晴 Nipponbare	均值±标准Mean±SD	变幅 Range	峰度Kurtosis	偏度Skewness
稻谷粒长	E1	9.86±0.47**	7.38±1.56	9.59±0.35	8.37~10.69	-0.46	2.58
Grain length / mm	E2	9.76±0.08**	7.68±0.16	9.31±0.28	8.23~9.77	-1.66	3.99
	E3	9.79±0.16**	7.67±0.19	9.52±0.33	8.05~10.14	-1.79	5.19
	E4	9.80±0.24**	7.63±0.29	9.78±0.31	8.71~10.30	-1.06	1.42
稻谷粒宽	E1	2.79±0.12*	3.24±0.60	2.78±0.13	2.55~3.53	2.33	10.17
Grain width / mm	E2	2.48±0.12*	3.30±0.56	2.99±0.12	2.69~3.61	2.00	8.05
	E3	2.59±0.19*	3.17±0.00	2.81±0.13	2.43~3.38	1.83	6.11
	E4	2.65±0.14*	3.13±0.42	2.83±0.11	2.56~3.27	1.43	4.11
稻谷长宽比	E1	3.55±0.27**	2.29±0.13	3.49±0.19	2.62~3.78	-1.91	4.62
Length-width ratio	E2	3.47±0.12**	2.31±0.23	3.14±0.16	2.42~3.38	1.60	3.70
	E3	3.78±0.11**	2.42±0.19	3.41±0.19	2.71~3.71	-1.59	2.73
	E4	3.70±0.05**	2.34±0.14	3.47±0.18	2.60~3.80	-1.82	5.04
稻谷粒厚	E1	2.26±0.09	2.17±0.05	2.19±0.05	2.04~2.34	0.07	0.64
Grain thickness / mm	E2	2.14±0.10	2.09±0.06	2.18±0.05	2.05~2.33	0.20	0.12
	E3	2.09±0.05	2.19±0.09	2.12±0.06	1.93~2.24	-0.54	0.05
	E4	1.99±0.06	2.01±0.12	2.10±0.05	1.95~2.20	-0.33	0.05
稻谷千粒重	E1	31.02±2.37**	22.44±1.69	29.51±2.29	22.17~34.21	-0.64	0.70
1000-grain weight / g	E2	31.36±3.06**	22.26±2.01	29.63±1.97	23.24~34.90	-0.40	0.77
	E3	30.39±2.74**	21.86±1.49	27.39±2.42	19.71~33.65	-0.21	0.48
	E4	32.53±2.36**	22.65±1.84	29.46±2.16	23.46~33.49	-0.58	0.41
糙米长	E1	6.87±0.31**	4.88±0.36	6.91±0.23	6.14~7.30	-1.50	2.67
Brown rice length	E2	6.84±0.41**	0.86±0.49	6.84±0.24	5.94~7.36	-1.64	3.57
/mm	E3	6.91±0.46**	4.99±0.61	6.93±0.24	5.96~7.39	-1.54	3.65
	E4	6.91±0.37**	4.70±0.23	6.92±0.27	5.97~7.54	-0.57	0.55
糙米宽	E1	2.38±0.12*	2.70±0.08	2.36±0.11	2.17~2.92	2.04	7.85
Brown rice width	E2	2.27±0.14*	2.67±0.13	2.52±0.09	2.31~3.01	1.90	9.25
/mm	E3	2.32±0.19*	2.78±0.05	2.40±0.09	2.16~2.79	1.50	5.41
	E4	2.46±0.11*	2.75±0.06	2.36±0.11	2.07~2.89	1.17	4.67
糙米长宽比	E1	2.98±0.21**	1.87±0.19	2.95±0.17	2.17~3.21	-1.67	3.84
Length-to-width ratio	E2	2.94±0.23**	1.84±0.17	2.72±0.14	2.11~2.99	-1.56	4.07
of brown rice	E3	2.98±0.31**	1.81±0.16	2.91±0.15	2.33~3.15	-1.53	3.09
	E4	2.76±0.27**	1.86±0.19	2.95±0.14	2.26~3.16	-1.95	5.56
糙米厚	E1	1.93±0.10	2.07±0.06*	1.93±0.05	1.79~2.10	0.14	0.66
Thickness of brown rice	E2	1.90±0.01	2.12±0.04*	1.91±0.05	1.77~2.08	0.36	1.48
/ mm	E3	1.85±0.07	1.96±0.06*	1.87±0.05	1.73~2.07	0.10	2.63
	E4	1.91±0.06	2.10±0.05*	1.88±0.05	1.77~2.06	0.37	0.38
糙米千粒重	E1	23.55±0.49*	21.88±0.32	24.64±1.57	20.47~29.46	0.03	0.33
1000-grain weight	E2	22.86±0.09*	21.76±0.36	24.32±1.37	20.34~28.20	-0.21	0.72
of brown rice / g	E3	22.29±0.14*	20.35±0.60	23.14±1.61	18.19~27.60	-0.10	0.31
	E4	23.67±0.52*	21.66±0.54	2.83±1.56	19.63~26.95	-0.39	0.00

QTL	区间 Interval			E1						E2						E3						E4
QTL	区间 Interval			LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%
qGL2	M123-M124																					3.32		-0.29		7.63
qGL3.1	M204-M205			6.61		-0.48		17.64
qGL3.2	M217-M218															3.40		-0.30		11.16
qGL3.3																						6.90		-0.43		17.02
qGL4.1	M245-M246			3.01		-0.38		7.48														3.04		-0.34		6.94
qGL4.2	M255-M256									3.90		-0.29		14.13
qGW1	M9-M10																					3.25		-0.07		5.10
qGW2.1	M142-M143									10.44		0.10		15.05
qGW2.2	M157-M158															4.56		0.13		9.17
qGW5.2	M334-M335															8.27		0.10		18.13
qGW5.1	M328-M329			13.14		0.14		35.23		21.04		0.16		38.10		12.77		0.12		30.29		22.19		0.20		51.01
qGW7	M491-M492			8.05		0.11		19.41
qGW8	M537-M538															4.64		-0.10		9.28
qGW9	M553-M554									8.72		-0.11		12.15
qGW10	M639-M640																					3.89		0.10		6.24
qLW2.1	M136-M137															4.28		-0.11		9.86
qLW2.2	M138-M139			5.24		-0.10		9.74
qLW3.2	M204-M205															3.47		-0.15		7.86
qLW3.1	M217-M218			10.41		-0.23		21.53
qLW3.3	M220-M221																					6.21		-0.21		16.81
qLW4.1	M245-M246			3.04		-0.16		5.40
qLW4.2	M255-M256									4.31		-0.14		13.34
qLW4.3	M276-M277															3.14		-0.10		7.07
qLW5	M328-M329			10.89		-0.18		22.75		6.25		-0.14		20.13		8.54		-0.17		21.5		8.92		-0.17		25.51
qLW7	M491-M492																					4.11		-0.12		10.67
qLW10.0	M643-M644			3.58		-0.12		6.44								4.42		-0.15		10.21
qGT1.1	M89-M90															6.42		-0.05		14.72
qGT1.2	M90-M91																					3.52		-0.04		10.40
qGT4	M260-M261															3.08		-0.04		6.59
qGT7.1	M488-M489			3.18		0.06		10.01
qGT7.2	M483-M484									3.31		0.05		12.42
qGT8.1	M540-M541			3.23		-0.04		10.17
qGT8.2	M537-M538															7.02		-0.09		16.30
qGT8.3	M539-M540																					4.89		-0.04		15.12
qTGW2	M153-M154															3.18		1.83		9.99
qTGW3	M204-M205			4.90		-2.66		12.58														3.11		-2.16		9.67
qTGW4.1	M299-M300			3.29		-1.51		7.96
qTGW4.2	M308-M309															3.02		-1.98		9.66
qTGW5.1	M363-M364									13.76		-10.14		11.55
qTGW5.2	M374-M375									22.19		9.6		30.05
qTGW8.2	M537-M538															3.29		-2.38		10.36		4.00		-2.47		12.68
qTGW8.1	M540-M541			4.47		-1.69		11.37
qBRL1.1	M61-M62															4.97		-0.23		7.75
qBRL1.2	M84-M85															3.08		0.13		4.62
qBRL2	M123-M124															4.82		-0.22		7.49
qBRL3.3	M193-M194															5.86		-0.18		9.30
qBRL3.1	M204-M205			3.84		-0.27		12.54
QTL		区间 Interval			E1						E2						E3						E4
QTL		区间 Interval			LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%
qBRL3.2			M217-M218								12.22		-0.38		30.96		9.17		-0.28		15.59
qBRL4.1			M245-M246								3.94		-0.28		8.40		4.70		-0.27		7.29		4.02		-0.36		10.74
qBRL5.1			M336-M337																				4.18		-0.20		11.41
qBRL8.1			M537-M538														4.07		-0.20		6.23
qBRL8.2			M549-M550																				3.66		-0.24		9.72
qBRW2			M143-M144														6.55		0.06		12.17
qBRW4			M228-M229																				3.44		-0.08		10.98
qBRW5.2			M328-M329														16.82		0.11		38.80
qBRW5.1			M324-M325		5.62		0.10		19.85		15.31		0.12		43.56
qBRW7			M488-M489								4.60		0.10		10.46								5.16		0.16		17.04
qBRW8.2			M537-M538														4.39		-0.07		7.82
qBRLW2.2			M136-M137																				3.31		-0.08		10.38
qBRLW2.1			M116-M117								3.33		-0.08		6.32
qBRLW3.1			M217-M218								12.09		-0.19		27.46
qBRLW3.2			M220-M221																				6.22		-0.18		20.65
qBRLW5			M324-M325		3.22		-0.12		11.89		9.64		-0.14		20.80
qBRLW7.1			M489-M490														5.23		-0.19		19.91
qBRLW7.2			M488-M489																				4.71		-0.19		15.17
qBRT1			M66-M67														5.70		-0.05		12.94
qBRT2			M170-M171														4.45		-0.04		10.26
qBRT7.2			M485-M486														5.09		0.05		11.41		4.33		0.06		13.69
qBRT7.1			M488-M489								4.59		0.07		16.39
qBRT8			M544-M545														4.98		-0.05		11.25
qBRTW1			M66-M67														3.84		-1.35		9.09
qBRTW2			M153-M154														3.02		1.07		7.01
qBRTW3			M204-M205								4.70		-1.41		9.58								3.16		-1.54		9.90
qBRTW5.1			M324-M325								6.81		1.24		14.48
qBRTW8.2			M537-M538														5.36		-1.86		13.03
qBRTW8.1			M539-M540								4.27		-1.17		8.79								4.77		-1.38		15.64
qBRTW11.1			M652-M653								4.84		-0.72		9.89

QTL	区间 Interval			E1						E2						E3						E4
QTL	区间 Interval			LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%
qGL2	M123-M124																					3.32		-0.29		7.63
qGL3.1	M204-M205			6.61		-0.48		17.64
qGL3.2	M217-M218															3.40		-0.30		11.16
qGL3.3																						6.90		-0.43		17.02
qGL4.1	M245-M246			3.01		-0.38		7.48														3.04		-0.34		6.94
qGL4.2	M255-M256									3.90		-0.29		14.13
qGW1	M9-M10																					3.25		-0.07		5.10
qGW2.1	M142-M143									10.44		0.10		15.05
qGW2.2	M157-M158															4.56		0.13		9.17
qGW5.2	M334-M335															8.27		0.10		18.13
qGW5.1	M328-M329			13.14		0.14		35.23		21.04		0.16		38.10		12.77		0.12		30.29		22.19		0.20		51.01
qGW7	M491-M492			8.05		0.11		19.41
qGW8	M537-M538															4.64		-0.10		9.28
qGW9	M553-M554									8.72		-0.11		12.15
qGW10	M639-M640																					3.89		0.10		6.24
qLW2.1	M136-M137															4.28		-0.11		9.86
qLW2.2	M138-M139			5.24		-0.10		9.74
qLW3.2	M204-M205															3.47		-0.15		7.86
qLW3.1	M217-M218			10.41		-0.23		21.53
qLW3.3	M220-M221																					6.21		-0.21		16.81
qLW4.1	M245-M246			3.04		-0.16		5.40
qLW4.2	M255-M256									4.31		-0.14		13.34
qLW4.3	M276-M277															3.14		-0.10		7.07
qLW5	M328-M329			10.89		-0.18		22.75		6.25		-0.14		20.13		8.54		-0.17		21.5		8.92		-0.17		25.51
qLW7	M491-M492																					4.11		-0.12		10.67
qLW10.0	M643-M644			3.58		-0.12		6.44								4.42		-0.15		10.21
qGT1.1	M89-M90															6.42		-0.05		14.72
qGT1.2	M90-M91																					3.52		-0.04		10.40
qGT4	M260-M261															3.08		-0.04		6.59
qGT7.1	M488-M489			3.18		0.06		10.01
qGT7.2	M483-M484									3.31		0.05		12.42
qGT8.1	M540-M541			3.23		-0.04		10.17
qGT8.2	M537-M538															7.02		-0.09		16.30
qGT8.3	M539-M540																					4.89		-0.04		15.12
qTGW2	M153-M154															3.18		1.83		9.99
qTGW3	M204-M205			4.90		-2.66		12.58														3.11		-2.16		9.67
qTGW4.1	M299-M300			3.29		-1.51		7.96
qTGW4.2	M308-M309															3.02		-1.98		9.66
qTGW5.1	M363-M364									13.76		-10.14		11.55
qTGW5.2	M374-M375									22.19		9.6		30.05
qTGW8.2	M537-M538															3.29		-2.38		10.36		4.00		-2.47		12.68
qTGW8.1	M540-M541			4.47		-1.69		11.37
qBRL1.1	M61-M62															4.97		-0.23		7.75
qBRL1.2	M84-M85															3.08		0.13		4.62
qBRL2	M123-M124															4.82		-0.22		7.49
qBRL3.3	M193-M194															5.86		-0.18		9.30
qBRL3.1	M204-M205			3.84		-0.27		12.54
QTL		区间 Interval			E1						E2						E3						E4
QTL		区间 Interval			LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%		LOD		加性效应Additive		贡献率PVE/%
qBRL3.2			M217-M218								12.22		-0.38		30.96		9.17		-0.28		15.59
qBRL4.1			M245-M246								3.94		-0.28		8.40		4.70		-0.27		7.29		4.02		-0.36		10.74
qBRL5.1			M336-M337																				4.18		-0.20		11.41
qBRL8.1			M537-M538														4.07		-0.20		6.23
qBRL8.2			M549-M550																				3.66		-0.24		9.72
qBRW2			M143-M144														6.55		0.06		12.17
qBRW4			M228-M229																				3.44		-0.08		10.98
qBRW5.2			M328-M329														16.82		0.11		38.80
qBRW5.1			M324-M325		5.62		0.10		19.85		15.31		0.12		43.56
qBRW7			M488-M489								4.60		0.10		10.46								5.16		0.16		17.04
qBRW8.2			M537-M538														4.39		-0.07		7.82
qBRLW2.2			M136-M137																				3.31		-0.08		10.38
qBRLW2.1			M116-M117								3.33		-0.08		6.32
qBRLW3.1			M217-M218								12.09		-0.19		27.46
qBRLW3.2			M220-M221																				6.22		-0.18		20.65
qBRLW5			M324-M325		3.22		-0.12		11.89		9.64		-0.14		20.80
qBRLW7.1			M489-M490														5.23		-0.19		19.91
qBRLW7.2			M488-M489																				4.71		-0.19		15.17
qBRT1			M66-M67														5.70		-0.05		12.94
qBRT2			M170-M171														4.45		-0.04		10.26
qBRT7.2			M485-M486														5.09		0.05		11.41		4.33		0.06		13.69
qBRT7.1			M488-M489								4.59		0.07		16.39
qBRT8			M544-M545														4.98		-0.05		11.25
qBRTW1			M66-M67														3.84		-1.35		9.09
qBRTW2			M153-M154														3.02		1.07		7.01
qBRTW3			M204-M205								4.70		-1.41		9.58								3.16		-1.54		9.90
qBRTW5.1			M324-M325								6.81		1.24		14.48
qBRTW8.2			M537-M538														5.36		-1.86		13.03
qBRTW8.1			M539-M540								4.27		-1.17		8.79								4.77		-1.38		15.64
qBRTW11.1			M652-M653								4.84		-0.72		9.89

染色体 Chr.	区间 Marker interval	多效性QTL Pleiotropic QTL	已克隆基因 Cloned gene	参考文献 Reference
1	M66-M67	qBRTW1, qBRT1	PGL2	Seonghoe et al. ^[21]
2	M123-M124	qGL2, qBRL2	GW2	Song et al. ^[12]
	M136-M137	qLW2.2, qBRLW2.2	GS2	Hu et al. ^[22]
	M153-M154	qTGW2, qBRTW2
3	M204-M205	qGL3.1, qLW3.2, qTGW3, qBRTW3, qBRL3.1	GS3	Liu et al. ^[13]
	M217-M218	qGL3.2, qLW3.1, qBRL3.2, qBRLW3.1
	M220-M221	qGL3.3, qLW3.3, qBRLW3.2	PGL1	Dany H et al. ^[23]
4	M245-M246	qGL4.1, qLW4.1, qBRL4.1	GL4	Wu et al. ^[24]
	M255-M256	qGL4.2, qLW4.2
5	M328-M329	qGW5.1, qLW5, qBRW5.2	gw5	Wan et al. ^[18]
	M324-M325	qBRW5.1, qBRLW5, qBRTW5.1
7	M491-M492	qGW7, qLW7	GW7	Wang et al. ^[25]
	M488-M489	qGT7.1, qBRW7, qBRLW7.2, qBRT7.1
8	M537-M538	qGW8, qGT8.2, qBRL8.1, qBRW8.2, qTGW8.2, qBRTW8.2	GW8	Wang et al. ^[26]
	M539-M540	qBRTW8.1, qGT8.3
	M540-M541	qTGW8.1, qGT8.1