四倍体小麦胚大小性状QTL定位与分析

doi:10.3864/j.issn.0578-1752.2023.02.001

Abstract

Abstract:

【Objective】 This study is to excavate embryo-related quantitative trait loci (QTL) with potential breeding value, to explore the genetic relationship between embryo and other agronomic traits in tetraploid wheat, and finally to aim at laying an important foundation for the fine mapping and breeding utilization of embryo-related traits in the future. 【Method】A total of 121 F₈ recombinant inbred lines (RIL) constructed by crossing tetraploid durum wheat (Ailanmai) and wild emmer wheat (LM001) were used. This RIL population was planted in five different environments including Chongzhou (2018-2020), Wenjiang (2020), and Ya'an (2020) in Sichuan Province for phenotypic evaluation of embryo length (EL), embryo width (EW), embryo length/embryo width (EL/EW), embryo length/kernel length (EL/KL), embryo width/kernel width (EW/KW), and embryo area (EA). QTL mapping was performed based on a genetic linkage map constructed based on the wheat 55K SNP. 【Result】 The embryo size-related traits showed approximately normal distribution in the RIL population satisfying the genetic characteristics of quantitative traits. A total of 27 QTL for embryo size-related traits were detected in five environments over three years. Among them, seven ones controlling EL could contribute 7.75%-21.74% of phenotypic variation. Seven QTLs controlling EW could explain 7.67%-33.29% of phenotypic variation. Five stable and major QTLs (QEL.sicau-AM-3B, QEW.sicau-AM-2B, QEW/KW.sicau-AM-2B, QEL/EW.sicau- AM-2B-1 and QEA.sicau-AM-2B) were identified, and they explained 11.88%-18.99%, 21.77%-29.41%, 8.80%-24.92%, 12.79%- 25.69% and 10.47%-15.22% of phenotypic variation, respectively. In addition, four QTL-rich regions were identified in the embryo size-related loci mentioned above. The QTL controlling EL/KL and EL was located on chromosome 1B, that for EW, EL/EW, EW/KW, and EA was located on 2B, that controlling EL and EA was on 3B, and that controlling EL/EW and EW/KW was on 6B. Embryo size was significantly and positively correlated with kernel size. Further, the major QTL for EL, QEL.sicau-AM-3B was co-located with that for kernel length identified previously, but that for EW QEW.sicau-AM-2B was independent of that for kernel width. Four genes likely involved in regulation of embryo size were identified in intervals where major QTL were mapped. 【Conclusion】Five stable and major QTLs were identified: QEL.sicau-AM-3B, QEW.sicau-AM-2B, QEW/KW.sicau-AM-2B, QEL/EW.sicau-AM-2B-1, QEA.sicau-AM-2B, among which QEW.sicau-AM-2B may be novel.

Key words: tetraploid wheat, embryo length, embryo width, embryo area, QTL mapping

CHEN JiHao, ZHOU JieGuang, QU XiangRu, WANG SuRong, TANG HuaPing, JIANG Yun, TANG LiWei, $\boxed{\hbox{LAN XiuJin}}$, WEI YuMing, ZHOU JingZhong, MA Jian. Mapping and Analysis of QTL for Embryo Size-Related Traits in Tetraploid Wheat[J].Scientia Agricultura Sinica, 2023, 56(2): 203-216.

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Figures/Tables 9

Table 1

Phenotypic variation of embryo related traits for parents and RIL in AM population"

性状 Trait	环境 Environment	亲本 Parents		重组自交系群体 RIL population
性状 Trait	环境 Environment	Ailanmai	LM001	范围 Range	平均值 Mean	标准差 SD	变异系数 CV	偏度 Skewness	峰度 Kurtosis	广义遗传力 H²
EL (mm)	2020WJ	2.58	3.09a	2.13—3.99	3.20	0.31	0.10	-0.38	0.81	0.75
	2020YA	2.55	3.08a	2.14—4.00	2.94	0.41	0.14	0.82	0.13
	2020CZ	2.51	3.08a	2.29—3.99	3.06	0.36	0.12	0.37	-0.22
	2019CZ	2.64	3.10a	2.13—4.00	3.07	0.35	0.11	0.12	0.01
	2018CZ	2.43	2.75a	2.43—4.00	3.13	0.31	0.10	0.21	0.04
	BLUP	2.62	3.02	2.45—3.37	3.00	0.17	0.06	0.59	0.81
EW (mm)	2020WJ	1.95a	1.88	1.48—2.95	2.02	0.33	0.16	0.84	0.05	0.76
	2020YA	2.14a	1.93	1.51—2.94	1.98	0.30	0.15	0.94	0.65
	2020CZ	1.98b	1.91	1.40—3.00	2.08	0.36	0.17	0.66	-0.12
	2019CZ	2.19a	1.99	1.43—3.00	2.06	0.32	0.16	1.03	1.19
	2018CZ	1.90a	1.54	1.45—3.00	2.01	0.30	0.15	1.16	1.48
	BLUP	2.02	1.86	1.64—3.00	1.95	0.14	0.07	0.14	0.01
EL/KL	2020WJ	0.39b	0.36	0.31—0.49	0.38	0.03	0.08	0.85	0.64	0.37
	2020YA	0.38b	0.36	0.27—0.41	0.34	0.03	0.09	0.12	-0.29
	2020CZ	0.40b	0.37	0.29—0.48	0.35	0.03	0.09	0.93	3.09
	2019CZ	0.40b	0.36	0.29—0.49	0.36	0.03	0.08	0.93	2.12
	2018CZ	0.39a	0.34	0.34—0.49	0.38	0.03	0.08	1.21	1.42
	BLUP	0.37	0.36	0.33—0.38	0.35	0.01	0.03	-0.66	1.88
EW/KW	2020WJ	0.50b	0.56	0.42—0.71	0.54	0.06	0.11	0.27	-0.29	0.64
	2020YA	0.56	0.57	0.44—0.81	0.56	0.07	0.13	0.76	0.76
	2020CZ	0.50b	0.57	0.39—0.76	0.57	0.07	0.12	0.18	0.23
	2019CZ	0.56	0.58	0.39—0.69	0.54	0.06	0.11	0.01	-0.15
	2018CZ	0.49b	0.46	0.42—0.65	0.55	0.06	0.11	0.26	-0.19
	BLUP	0.53	0.55	0.47—0.65	0.55	0.03	0.05	0.22	0.14
EL/EW	2020WJ	1.33b	1.65	1.20—1.99	1.65	0.16	0.10	-0.41	0.16	0.72
	2020YA	1.20	1.60a	1.15—1.99	1.49	0.16	0.11	0.71	0.68
	2020CZ	1.28	1.62a	1.12—2.52	1.54	0.2	0.13	1.78	7.93
	2019CZ	1.23	1.56	1.16—2.14	1.53	0.17	0.11	0.37	0.92
	2018CZ	1.28b	1.79	1.28—2.16	1.6	0.16	0.10	0.33	0.58
	BLUP	1.31	1.63	1.20—1.78	1.55	0.11	0.07	-0.52	0.47
EA (mm²)	2020WJ	3.62a	4.17	2.44—6.26	4.39	0.69	0.16	0.10	-0.01	0.75
	2020YA	3.93a	4.29	2.56—6.35	4.07	0.81	0.20	0.59	-0.22
	2020CZ	3.56a	4.24	2.65—6.18	4.27	0.72	0.17	-0.05	-0.16
	2019CZ	4.16a	4.44	2.74—6.17	4.35	0.73	0.17	0.25	-0.38
	2018CZ	3.33b	3.04	2.76—6.63	4.33	0.71	0.16	0.39	0.32
	BLUP	3.80	4.07	3.22—5.48	4.22	0.46	0.11	-0.12	-0.21

Table 1

Table 2

Fig. 1

Table 3

Table 4

QTL mapping results of embryo-related traits"

性状 Trait	数量性状位点 QTL	环境Environment	染色体 Chromosome	左标记 Left marker	右标记 Right marker	位置 Position (cM)	阈值 LOD	表型变异率 PVE (%)	加性效应 Add
胚长 EL	QEL.sicau-AM-3B	2020WJ	3B	AX-110375013	AX-110569582	6.91—7.34	5.15	16.21	-0.11
		2020YA	3B	AX-110375013	AX-110569582	6.91—7.34	3.63	11.88	-0.10
		2020CZ	3B	AX-111112626	AX-110375013	6.91—7.34	4.60	14.95	-0.10
		2019CZ	3B	AX-110375013	AX-110569582	6.91—7.34	4.90	18.99	-0.12
		BLUP	3B	AX-110375013	AX-110569582	6.91—7.34	8.41	21.74	-0.09
	QEL.sicau-AM-1B-1	2020CZ	1B	AX-108913437	AX-111510846	46.84—47.27	2.78	8.28	0.08
	QEL.sicau-AM-1B-1	BLUP	1B	AX-108913437	AX-111510846	46.84—47.27	4.22	9.77	0.06
	QEL.sicau-AM-1B-2	2020YA	1B	AX-111720572	AX-110579523	50.30—51.15	4.14	13.37	0.11
	QEL.sicau-AM-7B	2020WJ	7B	AX-110995675	AX-109951040	171.56—172.01	2.61	7.75	0.07
胚宽 EW	QEW.sicau-AM-2B	2020WJ	2B	AX-109390001	AX-110571221	0.00—1.92	7.42	24.13	0.10
		2020CZ	2B	AX-109390001	AX-110571221	0.00—1.92	7.41	22.40	0.10
		2019CZ	2B	AX-109390001	AX-110571221	0.00—1.92	6.19	21.77	0.09
		2018CZ	2B	AX-109390001	AX-110571221	0.00—1.92	6.50	29.41	0.11
		BLUP	2B	AX-109390001	AX-110571221	0.00—1.92	12.89	33.29	0.08
	QEW.sicau-AM-2A-1	2020WJ	2A	AX-109455008	AX-109024104	70.71—74.53	2.55	7.67	0.06
		2019CZ	2A	AX-109455008	AX-109024104	70.71—74.53	2.75	9.12	0.06
		BLUP	2A	AX-109455008	AX-109024104	70.71—74.53	4.80	10.74	0.05
	QEW.sicau-AM-2A-2	2020CZ	2A	AX-109934383	AX-109314401	98.92—105.82	3.31	9.79	0.07
胚长/粒长 EL/KL	QEL/KL.sicau-AM-1B-1	2020YA	1B	AX-109379021	AX-111720572	49.00—50.30	2.78	12.45	0.01
	QEL/KL.sicau-AM-1B-2	2020CZ	1B	AX-108913437	AX-111510846	46.84—47.27	3.07	12.68	0.01
	QEL/KL.sicau-AM-4B	2019CZ	4B	AX-109052054	AX-109931786	11.56—12.43	2.77	11.29	0.01
	QEL/KL.sicau-AM-5B	2019CZ	5B	AX-109990234	AX-108886889	76.90—79.56	2.54	10.13	-0.01
	QEL/KL.sicau-AM-2A	BLUP	2A	AX-109455008	AX-109024104	70.71—74.53	3.70	9.03	0.01
	QEL/KL.sicau-AM-2B	BLUP	2B	AX-109390001	AX-110571221	0.00—1.92	6.66	20.67	0.01
胚宽/粒宽 EW/KW	QEW/KW.sicau-AM-2B	2020CZ	2B	AX-109390001	AX-110571221	0.00—1.92	4.05	12.49	0.02
		2019CZ	2B	AX-109390001	AX-110571221	0.00—1.92	2.57	8.80	0.02
		2018CZ	2B	AX-109390001	AX-110571221	0.00—1.92	5.16	24.92	0.02
		BLUP	2B	AX-109390001	AX-110571221	0.00—1.92	6.65	20.68	0.01
	QEW/KW.sicau-AM-4A	2019CZ	4A	AX-109272723	AX-110524110	11.88—31.77	2.78	13.49	-0.02
	QEW/KW.sicau-AM-6B	2020CZ	6B	AX-111012548	AX-108984460	136.24—162.46	3.47	10.94	-0.02
胚长/胚宽 EL/EW	QEL/EW.sicau-AM-2B-1	2020WJ	2B	AX-109390001	AX-110571221	0.00—1.92	8.00	21.55	-0.08
		2020YA	2B	AX-109390001	AX-110571221	0.00—1.92	4.04	16.41	-0.09
		2020CZ	2B	AX-109390001	AX-110571221	0.00—1.92	8.83	25.69	-0.08
		2019CZ	2B	AX-109390001	AX-110571221	0.00—1.92	4.19	12.79	-0.06
		2018CZ	2B	AX-109390001	AX-110571221	0.00—1.92	5.45	23.41	-0.07
		BLUP	2B	AX-109390001	AX-110571221	0.00—1.92	11.64	31.13	-0.06
性状 Trait	数量性状位点 QTL	环境Environment	染色体 Chromosome	左标记 Left marker	右标记 Right marker	位置 Position (cM)	阈值 LOD	表型变异率 PVE (%)	加性效应 Add
	QEL/EW.sicau-AM-2B-2	2018CZ	2B	AX-111525915	AX-109297362	64.00—66.68	3.01	12.12	-0.05
	QEL/EW.sicau-AM-2B-3	2020WJ	2B	AX-110447950	AX-108942420	33.21—39.07	3.03	7.89	-0.05
	QEL/EW.sicau-AM-6B	2020CZ	6B	AX-111012548	AX-108984460	136.24—162.46	3.38	9.10	0.05
	QEL/EW.sicau-AM-2A-1	2020WJ	2A	AX-109314401	AX-110069674	105.82—124.56	3.14	8.05	-0.05
	QEL/EW.sicau-AM-2A-2	BLUP	2A	AX-110492886	AX-109523500	92.49—95.29	2.76	6.33	-0.03
胚面积 EA	QEA.sicau-AM-2B	2020WJ	2B	AX-109390001	AX-110571221	0.00—1.92	4.23	11.34	0.25
		2019CZ	2B	AX-109390001	AX-110571221	0.00—1.92	2.88	10.47	0.21
		2018CZ	2B	AX-109390001	AX-110571221	0.00—1.92	3.01	15.22	0.25
		BLUP	2B	AX-109390001	AX-110571221	0.00—1.92	6.66	20.67	0.01
	QEA.sicau-AM-2A-1	2020WJ	2A	AX-109437513	AX-109974497	66.15—67.06	4.29	11.41	0.25
	QEA.sicau-AM-2A-2	BLUP	2A	AX-109455008	AX-109024104	70.71—74.53	3.07	9.03	0.01
	QEA.sicau-AM-3B	2019CZ	3B	AX-110375013	AX-110569582	6.91—7.34	4.65	17.65	-0.28
	QEA.sicau-AM-4B	2020CZ	4B	AX-108737791	AX-110544397	20.75—22.08	3.47	13.18	0.23

Table 4

Fig. 2

Fig. 3

Table 5

Table 6

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性状 Trait	环境 Environment	2020WJ	2020YA	2020CZ	2019CZ	2018CZ	BLUP
胚长 EL	2020WJ	1
	2020YA	0.34**	1
	2020CZ	0.49**	0.38**	1
	2019CZ	0.37**	0.32**	0.48**	1
	2018CZ	0.35**	0.15**	0.39**	0.36**	1
	BLUP	0.69**	0.58**	0.68**	0.68**	0.65**	1
胚宽 EW	2020WJ	1
	2020YA	0.39**	1
	2020CZ	0.38**	0.38**	1
	2019CZ	0.43**	0.25**	0.47**	1
	2018CZ	0.36**	0.41	0.34**	0.26*	1
	BLUP	0.63**	0.66**	0.65**	0.60**	0.59**	1
胚长/粒长 EL/KL	2020WJ	1
	2020YA	0.06	1
	2020CZ	0.46**	0.19	1
	2019CZ	0.45**	0.08	0.43**	1
	2018CZ	0.37**	0.10	0.27*	0.31**	1
	BLUP	0.68**	0.39**	0.64**	0.62**	0.51**	1
胚宽/粒宽 EW/KW	2020WJ	1
	2020YA	0.29**	1
	2020CZ	0.36**	0.30**	1
	2019CZ	0.39**	0.35**	0.32**	1
	2018CZ	0.54**	0.35**	0.42**	0.32**	1
	BLUP	0.72**	0.69**	0.72**	0.68**	0.69**	1
胚长/胚宽 EL/EW	2020WJ	1
	2020YA	0.42**	1
	2020CZ	0.48**	0.61**	1
	2019CZ	0.46**	0.42**	0.56**	1
	2018CZ	0.63**	0.44**	0.43**	0.37**	1
	BLUP	0.78**	0.78**	0.73**	0.71**	0.73**	1
胚面积 EA	2020WJ	1
	2020YA	0.44**	1
	2020CZ	0.62**	0.37**	1
	2019CZ	0.58**	0.32**	0.48**	1
	2018CZ	0.51**	0.41**	0.56**	0.54**	1
	BLUP	0.80**	0.69**	0.79**	0.75**	0.75**	1

性状 Trait	QTL	遗传标记区间 Flanking markers	野生二粒小麦参考基因组 v2.0物理位置 Wild Emmer v2.0 physical map (Mb)	中国春参考基因组v2.1物理位置 Chines Spring wheat v2.1 physical map (Mb)
胚长 EL	QEL.sicau-AM-3B	AX-110375013—AX-110569582	695.42—696.83	690.52—692.06
胚长 EL	QEW.sicau-AM-2B	AX-109390001—AX-110571221	6.41—53.55	3.69—9.15
胚宽 EW	QEL/KL.sicau-AM-1B-1	AX-108913437—AX-111510846	409.17—411.07	395.61—397.54
胚宽 EW	QEL.sicau-AM-1B-2	AX-111720572—AX-110579523	432.54—447.52	418.79—434.50
胚长/粒长 EL/KL	QEW.sicau-AM-2A-1	AX-109455008—AX-109024104	614.32—618.45	615.12—619.25
胚宽/粒宽 EW/KW	QEW/KW.sicau-AM-2B	AX-109390001—AX-110571221	6.41—53.55	3.69—9.15
胚宽/粒宽 EW/KW	QEA.sicau-AM-2B	AX-109390001—AX-110571221	6.41—53.55	3.69—9.15
胚面积 EA	QEW/KW.sicau-AM-6B	AX-111012548—AX-108984460	712.45—723.67	721.64—730.61

性状 Trait	QTL	染色体Chromosome	左标记 Left marker		右标记 Right marker
性状 Trait	QTL	染色体Chromosome	标记ID Marker ID	物理位置 WE v2.0/CS 2.1 (Mb)	标记ID Marker ID	物理位置 WE v2.0/CS 2.1(Mb)
粒长Kernel length	qKL.3B	3B	*	*	Xwmc754	21.66/14.4
	QGl.ccsu-3B.1	3B	Xgwm566	91.51/77.7	Xgwm376	64.63/38.9
	QKL.sicau-AM-3B	3B	AX-110375013	695.42/690.52	AX-111112626	675.62/655.4
胚长EL	QEL.sicau-AM-3B	3B	AX-110375013	695.42/690.52	AX-110569582	696.83/692.06
胚宽EW	QEW.sicau-AM-2B	2B	AX-109390001	6.41/3.69	AX-110571221	53.55/9.15
千粒重 Thousand grain weight	QTgw.crc-2B	2B	Xwmc661	6.49/9.53	*	*
粒重Kernel weight	qKLW2B-1	2B	Xgwm501	98.15/680.09	Xmag3512	780.69/771.06
粒宽Kernel width	QKw.ncl-2B.1	2B	Xbarc183	123.83/2.60	Xbarc13	134.83/125.20
粒宽Kernel width	QTkw.ncl-2B.1	2B	Xbarc183	123.83/2.60	Xbarc7	134.82/125.20

Mapping and Analysis of QTL for Embryo Size-Related Traits in Tetraploid Wheat

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性状 Trait	分蘖数 Tiller number	株高 Plant height	粒长 Kernel length	粒宽 Kernel width	千粒重 Thousand-kernel weight	开花期 Flowering period
胚长 EL	0.03	0.62**	0.71**	0.33**	0.71**	-0.11
胚宽 EW	-0.06	0.16	0.02	0.36**	0.31**	0.08
胚长/粒长 EL/KL	-0.27**	0.08	-0.19*	0.36**	0.08	0.11
胚宽/粒宽 EW/KW	-0.12	-0.01	-0.07	-0.15	0.17	0.13
胚长/胚宽 EL/EW	0.07	0.33**	0.53**	-0.08	0.23*	-0.13
胚面积 EA	-0.02	0.42**	0.38**	0.40**	0.57**	-0.02

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