枣果实相关性状QTL定位分析

doi:10.16420/j.issn.0513-353x.2022-0056

摘要/Abstract

摘要：

以‘冬枣’×‘金丝4号’的103株F₁群体为试材，应用已构建的枣高密度遗传连锁图谱，采用复合区间作图法进行单果质量、果实纵径、横径、果形指数、可溶性固形物含量、总糖含量、总酸含量、糖酸比、维生素C含量、可食率等10个性状的QTL定位。10个性状在F₁代群体中表现为连续分布，具有数量性状的典型特征。共检测到157个与枣果实性状相关的QTL，其中单果质量相关44个，果实横径相关16个、纵径相关36个，果形指数相关9个，可溶性固形物相关11个，维生素C相关13个，总糖相关3个，总酸相关8个，糖酸比相关4个，可食率相关13个。各QTL位点的LOD值在2.65 ~ 6.25之间，可解释15.60% ~ 31.90%的表型变异。157个QTL分布在连锁图谱的10个连锁群中，其中5个连锁群上的27个位点同时连锁了2 ~ 3个表型，证实部分基因在对控制果实性状上存在一因多效。

关键词: 枣, 果实性状, 相关性, 数量性状位点

Abstract:

The separation of ten fruit characters（single fruit weight，fruit longitudinal diameter，fruit diameter，fruit shape index，soluble solids content，total sugar content，total acid content，sugar-acid ratio，vitamin C content，edible rate）in F₁ population of 103‘Dongzao’×‘Jinsi 4’and quantitative trait loci were analyzed by a high-density molecular genetic linkage map. Ten characters were distributed continuously on chromosomes，being typical quantitative characters. A total of 157 QTLs related to ten fruit traits were detected. Among these QTLs，44 were associated with the fruit weight，16 related to fruit diameter，36 associated with fruit longitudinal diameter and nine related to fruit shape index. The other QTLs were related to fruit quality，including 11 QTLs related to soluble solids，13 QTLs related to vitamin C，three QTLs related to total sugar，eight QTLs related to total acid and four QTLs related to sugar-acid ratio. Besides，there were 13 QTLs related to edible rate. The range of LOD for each QTL was 2.65-6.25，which can explain 15.60%-31.90% of phenotypic variation. One hundred and fifty-seven QTLs were distributed in ten linkage groups of linkage map，among which，27 loci in four linkage groups were linked with two to three phenotypes at the same time. The overlap of QTL mapping for different phenotypes suggested that some genes may be pleiotropic on the control of fruit traits.

Key words: Chinese jujube, fruit character, correlation, quantitative trait locus

中图分类号:

S665.1

唐海霞, 裴广营, 张琼, 王中堂. 枣果实相关性状QTL定位分析[J]. 园艺学报, 2023, 50(4): 754-764.

TANG Haixia, PEI Guangying, ZHANG Qiong, WANG Zhongtang. Location and Analysis of Quantitative Trait Loci in Chinese Jujube Fruit[J]. Acta Horticulturae Sinica, 2023, 50(4): 754-764.

图/表 5

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性状 Trait	F值 F value	母本 Female parent	父本 Male parent	亲中值 Median of parents	变异范围 Range	变异系数/% Coefficient of variation
单果质量/g Single fruit weight	32.02^**	15.00	10.59	12.57	3.15 ~ 26.16	19.49
果实横径/mm Fruit diameter	28.34^**	31.17	25.12	28.15	16.28 ~ 38.32	7.59
果实纵径/mm Fruit longitudinal diameter	44.27^**	30.34	32.75	31.55	17.24 ~ 41.38	7.22
果形指数Fruit shape index	1.72^**	0.97	1.30	1.14	0.81 ~ 1.42	7.91
可溶性固形物/% Soluble solid	12.30^**	27.71	29.35	28.53	15.90 ~ 41.50	8.64
可食率/% Edible rate	38.04^**	96.33	96.96	96.65	91.35 ~ 98.04	1.12
维生素C/（mg · g^-1）Vitamin C	608.34^**	2.25	4.16	3.21	1.49 ~ 4.26	24.93
总糖/（mg · g^-1）Total sugar	147.70^**	267.84	280.34	193.84	89.44 ~ 228.78	20.27
总酸/% Total acid	826.32^**	0.31	0.23	0.27	0.28 ~ 1.04	25.62
糖酸比 Suger-acid ratio	232.46	86.40	121.88	104.10	13.22 ~ 80.27	33.49

性状 Trait	F值 F value	母本 Female parent	父本 Male parent	亲中值 Median of parents	变异范围 Range	变异系数/% Coefficient of variation
单果质量/g Single fruit weight	32.02^**	15.00	10.59	12.57	3.15 ~ 26.16	19.49
果实横径/mm Fruit diameter	28.34^**	31.17	25.12	28.15	16.28 ~ 38.32	7.59
果实纵径/mm Fruit longitudinal diameter	44.27^**	30.34	32.75	31.55	17.24 ~ 41.38	7.22
果形指数Fruit shape index	1.72^**	0.97	1.30	1.14	0.81 ~ 1.42	7.91
可溶性固形物/% Soluble solid	12.30^**	27.71	29.35	28.53	15.90 ~ 41.50	8.64
可食率/% Edible rate	38.04^**	96.33	96.96	96.65	91.35 ~ 98.04	1.12
维生素C/（mg · g^-1）Vitamin C	608.34^**	2.25	4.16	3.21	1.49 ~ 4.26	24.93
总糖/（mg · g^-1）Total sugar	147.70^**	267.84	280.34	193.84	89.44 ~ 228.78	20.27
总酸/% Total acid	826.32^**	0.31	0.23	0.27	0.28 ~ 1.04	25.62
糖酸比 Suger-acid ratio	232.46	86.40	121.88	104.10	13.22 ~ 80.27	33.49

性状 Trait	最大值Maximum	最小值Minimum	平均值 Mean	正态性检验卡方值 Normal test chi-square values	P值 P value
单果质量/g Single fruit weight	26.16	3.15	10.24	6.33	0.61
横径/mm Fruit diameter	38.32	16.28	25.75	3.31	0.65
纵径/mm Fruit longitudinal diameter	41.38	17.24	27.51	2.24	0.82
果形指数Fruit shape index	1.42	0.81	1.08	5.15	0.40
可溶性固形物/% Soluble solids	41.50	15.90	29.80	42.15	< 0.05
可食率/% Edible rate	98.83	93.65	95.80	17.08	< 0.05
维生素C/（mg · g^-1）Vitamin C	4.26	1.49	2.23	8.71	0.12
总糖/（mg · g^-1）Total sugar	355.28	189.44	167.91	12.29	< 0.05
总酸/% Total acid	1.04	0.28	0.51	5.37	0.37
糖酸比Suger-acid ratio	80.27	13.22	62.60	3.04	0.69

性状 Trait	最大值Maximum	最小值Minimum	平均值 Mean	正态性检验卡方值 Normal test chi-square values	P值 P value
单果质量/g Single fruit weight	26.16	3.15	10.24	6.33	0.61
横径/mm Fruit diameter	38.32	16.28	25.75	3.31	0.65
纵径/mm Fruit longitudinal diameter	41.38	17.24	27.51	2.24	0.82
果形指数Fruit shape index	1.42	0.81	1.08	5.15	0.40
可溶性固形物/% Soluble solids	41.50	15.90	29.80	42.15	< 0.05
可食率/% Edible rate	98.83	93.65	95.80	17.08	< 0.05
维生素C/（mg · g^-1）Vitamin C	4.26	1.49	2.23	8.71	0.12
总糖/（mg · g^-1）Total sugar	355.28	189.44	167.91	12.29	< 0.05
总酸/% Total acid	1.04	0.28	0.51	5.37	0.37
糖酸比Suger-acid ratio	80.27	13.22	62.60	3.04	0.69

性状 Trait	性状位点 QTL		连锁群 Linkage group	区间范围/cM Confidence interval	峰值标记位置/cM Peak position	峰值标记 Marker			阈值 LOD	解释率/% Explain rate
单果质量 Single fruit weight	SFW1.1		lg1	4.37	4.37	chr1-606			3.37	18.70
	SFW1.2		lg1	5.39	5.39	chr1-499			3.15	17.60
	SFW1.3		lg1	8.37	8.37	chr1-569			3.44	19.10
	SFW1.4		lg1	8.51	8.51	chr1-485			3.44	19.10
	SFW1.5		lg1	9.74	9.74	chr1-497			3.15	17.60
	SFW1.6		lg1	12.63 ~ 14.34	14.34	chr1-695			3.37	18.70
	SFW1.7		lg1	15.84	15.84	chr1-829			3.03	17.00
	SFW1.8		lg1	19.15 ~ 21.52	21.16	chr1-953			4.43	23.80
	SFW1.9		lg1	28.19 ~ 28.59	28.51	unchr-12446			4.77	25.40
	SFW1.10		lg1	37.47	37.47	chr1-1060			3.61	19.90
	SFW1.11		lg1	37.83	37.83	chr1-1073			3.35	18.60
	SFW1.12		lg1	51.48	51.48	unchr-13300			3.05	17.10
	SFW4.1		lg4	8.76	8.76	chr4-4192			3.41	18.90
	SFW5.1		lg5	20.04 ~ 20.22	20.04	chr5-5151			3.62	19.90
	SFW6.1		lg6	22.46	22.46	chr5-5594			3.79	20.70
	SFW6.2		lg6	42.33	42.33	chr6-6345			3.02	16.90
	SFW11.1		lg11	43.73	43.73	chr11-10913			3.26	18.10
	SFW12.1		lg12	8.23 ~ 8.26	8.23	chr12-11184			3.23	18.00
	SFW12.2		lg12	11.21	11.21	chr6-6179			3.25	18.10
	SFW12.3		lg12	11.70	11.70	chr12-11222			3.14	17.50
	SFW12.4		lg12	13.05 ~ 13.06	13.05	chr12-11277			3.36	18.60
	SFW12.5		lg12	13.13	13.13	chr12-11433			3.77	20.70
	SFW12.6		lg12	13.17 ~ 13.33	13.20	chr12-11371			3.16	17.60
	SFW12.7		lg12	13.51 ~ 13.57	13.51	chr12-11415			3.48	19.20
	SFW12.8		lg12	13.77 ~ 13.79	13.79	chr12-11414			3.64	20.00
	SFW12.9		lg12	13.87	13.87	chr12-11418			3.28	18.30
	SFW12.10		lg12	14.20	14.20	unchr-12456			3.17	17.70
	SFW12.11		lg12	14.23	14.23	chr4-4277			3.32	18.40
	SFW12.12		lg12	14.62 ~ 14.63	14.62	unchr-12023			3.19	17.80
	SFW12.13		lg12	14.69 ~ 14.97	14.97	chr12-11360			3.43	19.00
	SFW12.14		lg12	15.08 ~ 15.09	15.08	unchr-12019			3.42	18.90
	SFW12.15		lg12	15.16	15.16	chr12-11404			3.04	17.00
	SFW12.16		lg12	15.59 ~ 15.73	15.73	chr12-11427			3.64	20.00
	SFW12.17		lg12	16.20	16.20	chr12-11443			3.75	20.60
	SFW12.18		lg12	16.45 ~ 17.79	16.45	chr12-11451			3.79	20.80
	SFW12.19		lg12	19.22	19.22	unchr-13403			3.08	17.20
	SFW12.20		lg12	19.84 ~ 22.71	22.01	chr12-11454			3.83	20.90
	SFW12.21		lg12	25.26 ~ 26.56	26.26	chr12-11500			4.16	22.60
	SFW12.22		lg12	29.17	29.17	unchr-12838			3.09	17.30
	SFW12.23		lg12	34.84 ~ 47.57	47.57	chr12-11669			5.09	26.80
	SFW12.24		lg12	47.75 ~ 50.47	47.76	chr12-11630			3.95	21.60
	SFW12.25		lg12	54.39 ~ 57.18	57.18	chr12-11737			3.76	20.60
	SFW12.26		lg12	61.73 ~ 62.62	61.73	unchr-12089			3.52	19.40
	SFW12.27		lg12	69.71	69.71	unchr-13124			3.43	19.00
果实横径	FD1.1		lg1	20.30 ~ 21.52	21.16	chr1-935			3.55	19.60
Fruit diameter	FD1.2		lg1	28.51 ~ 28.59	28.51	unchr-12446			4.15	22.50
	FD4.1		lg4	8.56	8.56	chr4-3856			3.09	17.30
	FD4.2		lg4	8.76	8.76	chr4-4192			3.22	17.90
	FD12.1		lg12	15.73	15.73	chr12-11427			3.23	18.00
	FD12.2		lg12	16.45 ~ 16.61	16.52	chr12-11449			3.86	21.10
	FD12.3		lg12	20.99 ~ 22.18	22.18	chr12-11455			3.28	18.20
	FD12.4		lg12	26.56 ~ 27.51	26.56	chr12-11509			3.16	17.40
	FD12.5		lg12	29.17	29.17	unchr-12838			3.16	17.40
	FD12.6		lg12	41.53 ~ 41.82	41.53	chr12-11559			3.24	18.10
	FD12.7		lg12	43.25	43.25	chr12-11564			3.02	16.90
	FD12.8		lg12	47.42 ~ 47.57	47.57	chr12-11669			4.11	22.30
	FD12.9		lg12	47.75 ~ 47.77	47.76	chr12-11630			3.59	19.80
	FD12.10		lg12	48.15 ~ 49.29	48.17	chr12-11646			3.44	19.00
	FD12.11		lg12	50.47	50.47	chr12-11591			3.56	19.60
果实纵径 Fruit longitudinal diameter	FD12.12		lg12	54.98	54.98		chr12-11734	3.10			17.30
	FL1.1		lg1	4.37	4.37		chr1-606	3.08			17.20
	FL1.2		lg1	19.35 ~ 21.52	21.16		chr1-935	4.17			22.60
	FL1.3		lg1	28.09 ~ 28.59	28.51		unchr-12446	5.19			27.30
	FL1.4		lg1	37.47	37.47		chr1-1060	3.08			17.20
	FL1.5		lg1	48.64 ~ 48.86	48.86		chr1-1184	3.15			17.60
	FL1.6		lg1	50.03	50.03		chr1-1189	3.14			17.60
	FL1.7		lg1	51.48	51.48		unchr-13300	3.12			17.40
	FL2.1		lg2	12.22	12.22		chr2-1579	3.01			16.90
	FL5.1		lg5	20.04 ~ 20.22	20.04		chr5-5151	3.46			19.20
	FL5.2		lg5	21.97	21.97		chr5-5211	3.02			16.90
	FL6.1		lg6	42.33	42.33		chr6-6345	3.43			19.00
	FL12.1		lg12	13.05 ~ 13.06	13.05		chr12-11277	3.24			18.00
	FL12.2		lg12	13.13	13.13		chr12-11433	3.56			19.60
	FL12.3		lg12	13.17 ~ 13.2	13.20		chr12-11371	3.22			17.90
	FL12.4		lg12	13.51 ~ 13.57	13.57		chr12-11421	3.23			18.00
果形指数 Fruit shape index	FL12.5		lg12	13.77 ~ 13.79	13.79		chr12-11414	3.87			21.10
	FL12.6		lg12	13.87	13.87		chr12-11418	3.28			18.20
	FL12.7		lg12	14.20	14.20		unchr-12456	3.36			18.60
	FL12.8		lg12	14.23	14.23		chr4-4277	3.07			17.20
	FL12.9		lg12	14.69 ~ 14.97	14.97		chr12-11360	3.97			21.60
	FL12.10		lg12	15.02 ~ 15.04	15.04		chr4-4272	3.36			18.70
	FL12.11		lg12	15.08 ~ 15.16	15.08		unchr-12019	3.96			21.60
	FL12.12		lg12	15.58 ~ 15.73	15.73		chr12-11427	3.41			18.90
	FL12.13		lg12	16.20	16.20		chr12-11443	3.52			19.40
	FL12.14		lg12	16.45 ~ 18.55	16.92		chr12-11377	3.71			20.40
	FL12.15		lg12	19.22 ~ 22.71	21.30		chr12-11477	3.97			21.60
	FL12.16		lg12	25.26 ~ 29.17	26.56		chr12-11509	3.41			18.90
	FL12.17		lg12	34.84 ~ 47.57	47.57		chr12-11669	6.25			31.90
	FL12.18		lg12	47.75 ~ 50.47	47.75		chr12-11643	4.48			24.10
	FL12.19		lg12	50.99 ~ 51.23	51.23		chr12-11713	3.27			18.20
	FL12.20		lg12	54.39 ~ 58.74	57.18		chr12-11737	4.81			25.60
	FL12.21		lg12	60.56 ~ 62.8	61.88		chr12-11800	4.56			24.40
	FL12.22		lg12	65.75	65.75		chr12-11858	3.18			17.70
	FL12.23		lg12	67.43 ~ 67.44	67.44		chr12-11880	3.33			18.50
	FL12.24		lg12	68.58 ~ 68.71	68.58		chr12-11865	3.35			18.60
	FL12.25		lg12	69.66 ~ 69.71	69.71		unchr-13124	3.59			19.80
	FS5.1		lg5	44.04 ~ 44.08	44.08		chr5-5512	5.81			30.00
	FS10.1		lg10	74.70 ~ 75.52	75.52		chr10-10116	3.78			20.70
	FS12.1		lg12	46.65 ~ 47.43	46.65		chr12-11706	3.43			19.00
	FS12.2		lg12	47.47	47.47		chr12-11616	3.01			16.90
	FS12.3		lg12	47.74	47.74		chr12-11584	3.12			17.40
	FS12.4		lg12	49.61	49.61		chr12-11601	3.11			17.40
	FS12.5		lg12	54.39 ~ 62.62	57.18		chr12-11737	4.08			22.20
	FS12.6		lg12	64.15	64.15		chr12-11755	3.26			18.10
	FS12.7		lg12	67.43 ~ 67.6	67.60		chr12-11903	3.20			17.80
可溶性固形物	SS1.1		lg1	29.66	29.66		chr1-100	3.24			18.00
Soluble solid	SS1.2		lg1	46.93	46.93		chr1-1163	3.31			18.40
	SS1.3		lg1	47.79 ~ 48.08	48.06		chr1-1186	3.82			20.90
	SS1.4		lg1	51.23 ~ 51.38	51.38		chr1-109	3.83			21.00
	SS1.5		lg1	51.48 ~ 51.98	51.69		chr1-105	4.04			22.00
	SS1.6		lg1	52.22	52.22		chr1-160	3.06			17.10
	SS1.7		lg1	52.47	52.47		chr1-157	3.18			17.70
	SS1.8		lg1	53.51	53.51		chr1-161	3.31			18.40
	SS1.9		lg1	55.17	55.17		chr1-91	3.04			17.10
	SS1.10		lg1	55.75	55.75		chr1-68	3.21			17.90
	SS11.1		lg11	10.32	10.32		chr11-10916	4.56			24.40
可食率	ER1.1		lg1	58.24	58.24		chr11-11137	3.45			19.10
Edible rate	ER6.1		lg6	29.56	29.56		unchr-12741	3.42			18.90
	ER6.2		lg6	31.40	31.40		chr6-6488	3.43			19.00
	ER6.3	lg6		32.12	32.12	chr6-6462			3.05	17.10
	ER6.4	lg6		34.22	34.22	chr6-6403			3.02	16.90
	ER9.1	lg9		0	0	chr9-8739			3.23	18.00
	ER9.2	lg9		12.34 ~ 16.99	13.31	chr9-8573			4.40	23.70
	ER9.3	lg9		17.65 ~ 24.97	24.97	chr9-8945			5.17	27.20
	ER9.4	lg9		27.61 ~ 29.67	27.61	chr9-8968			3.73	20.50
	ER9.5	lg9		31.48 ~ 34.08	31.48	unchr-13059			3.49	19.30
	ER12.1	lg12		9.64 ~ 11.54	11.21	chr6-6179			3.42	19.00
维生素C Vitamin C	ER12.2	lg12		30.98 ~ 45.54	33.00	chr12-11543			3.91	21.30
	ER12.3	lg12		47.06 ~ 50.47	47.43	chr12-11603			3.83	21.00
	VC7.1	lg7		59.91 ~ 60.09	60.09	chr7-7552			4.39	24.50
	VC9.1	lg9		4.91 ~ 5.46	5.37	chr9-8771			3.20	18.50
	VC9.2	lg9		9.74	9.74	chr9-8608			3.15	18.20
	VC9.3	lg9		10.21 ~ 10.75	10.21	chr9-8582			3.54	20.30
	VC9.4	lg9		12.18	12.18	chr9-8552			3.12	18.10
	VC9.5	lg9		12.69 ~ 17.46	17.13	chr9-8928			4.24	23.80
	VC9.6	lg9		17.67 ~ 24.97	23.69	chr9-8930			4.52	25.10
	VC12.1	lg12		7.52	7.52	unchr-12881			3.13	18.10
	VC12.2	lg12		9.57	9.57	chr12-11281			3.17	18.30
	VC12.3	lg12		13.17	13.17	chr12-11433			3.30	19.00
	VC12.4	lg12		15.90	15.90	chr12-11367			3.60	20.60
	VC12.5	lg12		26.56 ~ 30.98	29.17	unchr-12838			4.93	27.00
	VC12.6	lg12		32.80 ~ 35.10	32.80	chr12-11537			3.40	19.50
总糖	TS5.1	lg5		44.34 ~ 44.35	44.34	chr5-5516			2.92	17.10
Total sugar	TS10.1	lg10		69.22	69.22	chr10-9975			2.65	15.60
	TS12.1	lg12		13.87	13.87	chr12-11418			2.69	15.80
总酸	TA7.1	lg7		56.69	56.69	chr7-7485			3.96	22.40
Total acid	TA7.2	lg7		37.21	67.21	chr7-7596			3.05	17.70
	TA7.3	lg7		67.74	67.74	chr7-7699			3.84	21.80
	TA10.1	lg10		35.47	35.47	chr10-10200			4.01	22.60
	TA12.1	lg12		14.59	14.59	chr11-11152			3.45	19.80
	TA12.2	lg12		15.45	15.45	chr12-11362			3.22	18.60
	TA12.3	lg12		25.26	25.26	chr12-11500			3.22	18.60
	TA12.4	lg12		29.17	29.17	unchr-12838			3.33	19.20
糖酸比	SAR9.1	lg9		34.89	34.89	chr2-2060			3.05	17.70
Sugar-acid ratio	SAR12.1	lg12		16.28	16.28	chr12-11483			3.55	20.30
	SAR12.2	lg12		22.71	22.71	chr12-11468			3.11	18.00
	SAR12.3	lg12		29.17	29.17	unchr-12838			3.73	21.20