春小麦籽粒主要品质性状的全基因组关联分析

doi:10.3864/j.issn.0578-1752.2021.19.001

摘要/Abstract

摘要：

【目的】挖掘小麦籽粒品质性状显著相关的SNP位点及候选基因,并揭示其遗传机理,为相关基因克隆和分子标记辅助选择提供理论依据。【方法】通过检测298份国内外春小麦品种（系）5个环境下蛋白质含量、湿面筋含量、沉降值、淀粉含量、籽粒硬度、出粉率和容重等7个籽粒品质性状的表型,并结合小麦55K SNP芯片,采用Q+K关联混合模型进行全基因组关联分析（genome-wide association study,GWAS）。【结果】外引品种（系）、地方品种（系）和育成品种（系）的7个品质性状在不同环境下的变异系数分别为1.3%—13.4%、1.1%—18.6%和1.0%—13.9%。其中,外引品种（系）的蛋白质含量、湿面筋含量和沉降值的变异系数均为最高;新疆自育品种的淀粉含量、籽粒硬度和出粉率的变异系数最大,而新疆地方品种的蛋白质含量、湿面筋含量、沉降值、淀粉含量、籽粒硬度和出粉率6个品质性状的变异系数均介于外引品种（系）和新疆自育品种（系）之间。群体结构分析表明,298份小麦品种（系）可分为3个亚群。其中,亚群1包含128份（43.0%）试验材料,主要是来自新疆的地方品种（系）;亚群2包含24份（8.1%）试验材料,主要包括外引品种（系）和新疆地方品种;亚群3包含146份（48.9%）试验材料,主要是外引品种（系）。连锁不平衡分析表明A、B和D基因组及全基因组的LD衰减距离分别为10、10、6和8 Mb,依据全基因组的LD衰减距离,将在物理图谱上前后8 Mb区间内的位点认定为一个候选位点。通过GWAS共检测到85个与7个小麦籽粒品质性状显著关联的稳定位点（P<0.001）贡献率为3.7%—10.9%。在1B、1D、2D、3A、3D、4A、4B、5A、6A、6D、7A和7D染色体上均检测到稳定且同时与多个性状关联的位点。其中,7A染色体上的AX-109452823—AX-110545157同时与蛋白质含量、淀粉含量、湿面筋含量、沉降值、出粉率和籽粒硬度相关,且同时在4个环境中均被检测到。对稳定的位点进行候选基因发掘,筛选到10个可能与小麦籽粒品质相关的候选基因。其中TraesCS4A01G299800（阳离子氨基酸转运蛋白）、TraesCS7A01G059500（色氨酸脱羧酶）、TraesCS7A01G331200和TraesCS7D01G418700（木葡聚糖内转葡糖基酶/水解酶）对调控小麦籽粒氨基酸含量有重要作用。【结论】检测到85个稳定的且与小麦籽粒品质性状关联的位点,并筛选出10个与小麦籽粒品质性状相关的候选基因。

关键词: 小麦, 籽粒品质性状, 全基因组关联分析, SNP, 候选基因

Abstract:

【Objective】Identify SNPs and candidate genes that are significantly related to wheat grain quality traits, and reveal their genetic mechanism. 【Method】In this study, 298 introduced varieties (lines), Xinjiang landrace (lines) and Xinjiang bred varieties (lines) were used for an association population. Seven grain quality traits, including protein content (PRC), wet gluten content (WGC), sedimentation value (SV), starch content (STC), grain hardness (GH), flour yield (FY) and test weight (TW), were measured under five environments. Based on phenotypes of seven quality traits and genotypes of 55K SNP markers in this population, the Q+K association mixed model was used for genome-wide association analysis to obtain significantly associated SNP loci.【Result】The coefficients of variation of the seven grain quality traits of introduced varieties (lines), landraces and bred varieties (lines) under different environments were 1.3%-13.4%, 1.1%-18.6% and 1.0%-13.9%, respectively. Among them, the protein content, wet gluten content and sedimentation value of introduced varieties (lines) have the highest coefficient of variation (CV); Xinjiang bred varieties (lines) have the largest CV of starch content, grain hardness and flour yield. Whereas, for other six grain quality traits, including protein content, wet gluten content, sedimentation value, starch content, grain hardness, and the coefficients of variation of Xinjiang landraces are all between those of the introduced varieties (lines) and Xinjiang bred varieties (lines). Population structure analysis showed that 298 wheat varieties (lines) can be divided into 3 subgroups. Subgroup 1 contains 128 (43.0%) the materials mainly from landrace (lines); Subgroup 2 has 24 (8.1%) materials, mainly including introduced varieties (lines) and landraces; Subgroup 3 contains 146 (48.9%) materials, mainly introduced varieties (lines). The linkage disequilibrium analysis showed that the LD attenuation distances of the A, B and D genomes and the whole genome respectively were 10, 10, 6 and 8 Mb, according to the LD attenuation distance of the whole genome, the loci in the 8 Mb interval after the physical map were identified as a candidate loci. A total of 85 loci were simultaneously detected in two or more environments, that were significantly associated with 7 wheat grain quality traits detected by GWAS, with a contribution rate of 3.7%-10.9%. Stable SNPs associated with multiple traits were detected on chromosomes 1B, 1D, 2D, 3A, 3D, 4A, 4B, 5A, 6A, 6D, 7A and 7D. Among them, AX-109452823-AX-110545157 on chromosome 7A is related to protein content, starch content, wet gluten content, sedimentation value, flour yield and grain hardness, and was detected across four environments. Candidate genes at stable loci associated with multiple traits were searched, and 10 candidate genes that might be related to wheat grain quality were screened. Among them, TraesCS4A01G299800 (cationic amino acid transporter), TraesCS7A01G059500 (tryptophan decarboxylase), TraesCS7A01G331200, TraesCS7D01G418700 (xyloglucan endoglucosylase/hydrolase) play important roles in regulating the amino acid content in wheat grains.【Conclusion】The 85 loci were simultaneously detected in two or more environments, and 10 candidate genes related to wheat grain quality traits were predicted.

Key words: wheat, quality traits, genome-wide association analysis, SNP, candidate genes

严勇亮,时晓磊,张金波,耿洪伟,肖菁,路子峰,倪中福,丛花. 春小麦籽粒主要品质性状的全基因组关联分析[J]. 中国农业科学, 2021, 54(19): 4033-4047.

YAN YongLiang,SHI XiaoLei,ZHANG JinBo,GENG HongWei,XIAO Jing,LU ZiFeng,NI ZhongFu,CONG Hua. Genome-Wide Association Study of Grain Quality Related Characteristics of Spring Wheat[J]. Scientia Agricultura Sinica, 2021, 54(19): 4033-4047.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2021.19.001

https://www.chinaagrisci.com/CN/Y2021/V54/I19/4033

图/表 7

表1

春小麦品质性状表型变异"

性状 Trait	环境 Environment	最大值 Max			最小值 Min			平均值 Mean			标准差 SD			变异系数 CV (%)
性状 Trait	环境 Environment	外引种 Introduced varieties	地方种 Xinjiang landrace	育成种 Xinjiang bred varieties	外引种 Introduced varieties	地方种 Xinjiang landrace	育成种 Xinjiang bred varieties	外引种 Introduced varieties	地方种 Xinjiang landrace	育成种 Xinjiang bred varieties	外引种 Introduced varieties	地方种 Xinjiang landrace	育成种 Xinjiang bred varieties	外引种 Introduced varieties	地方种 Xinjiang landrace	育成种 Xinjiang bred varieties
蛋白质含量 PRC (%)	E1	22.60	22.03	19.79	13.96	13.40	14.22	18.67	16.53	16.81	1.70	1.71	1.51	9.13	10.36	8.98
	E2	21.40	23.24	19.97	14.72	13.18	12.93	17.95	16.16	15.80	1.42	1.87	1.15	7.89	11.60	7.27
	E3	22.02	22.79	18.84	13.92	13.24	13.48	17.61	16.17	15.45	1.38	1.48	1.14	7.84	9.12	7.40
	E4	20.79	21.83	19.71	13.99	12.37	13.18	16.60	16.18	15.85	1.42	1.58	1.15	8.54	9.74	7.28
	E5	23.13	22.72	18.51	15.03	15.16	14.59	18.59	17.68	16.45	1.35	1.39	0.91	7.26	7.87	5.54
湿面筋含量 WGC (%)	E1	51.27	50.05	45.15	31.90	29.18	31.43	42.03	36.76	37.93	3.88	4.19	3.29	9.23	11.40	8.68
	E2	48.52	52.90	45.46	33.19	28.44	27.97	40.39	35.88	35.80	3.33	4.57	2.72	8.23	12.73	7.61
	E3	50.05	52.44	43.09	31.76	28.97	29.88	39.90	36.09	35.35	3.21	3.58	2.66	8.05	9.93	7.52
	E4	46.74	49.74	44.84	31.84	26.92	29.13	37.61	36.17	36.14	3.18	3.76	2.67	8.46	10.39	7.39
	E5	52.26	51.98	40.80	34.31	33.33	32.40	41.84	39.54	37.67	3.11	3.41	2.07	7.42	8.61	5.51
沉降值 SV (mL)	E1	62.27	63.12	51.69	31.06	26.64	30.23	46.69	39.78	39.17	6.00	6.40	4.82	12.86	16.08	12.31
	E2	58.03	60.47	45.38	30.57	23.30	25.38	42.61	35.93	34.05	4.74	6.68	3.63	11.13	18.58	10.67
	E3	56.52	59.74	42.38	27.60	22.37	23.75	38.62	34.86	30.38	5.19	5.90	4.22	13.44	16.93	13.88
	E4	50.56	57.78	43.56	25.21	18.66	25.09	38.54	35.72	32.19	4.83	6.40	3.98	12.52	17.93	12.37
	E5	56.48	64.21	47.12	26.42	29.08	28.06	38.32	39.66	35.16	4.90	5.36	3.63	12.78	13.51	10.32
淀粉含量 STC (%)	E1	72.62	72.24	72.87	61.06	57.76	65.01	66.35	68.71	69.51	2.46	2.35	1.51	3.71	3.42	2.17
	E2	72.35	72.92	73.93	61.90	61.81	65.98	68.22	70.07	70.78	1.89	1.74	1.32	2.77	2.49	1.86
	E3	71.73	71.50	72.66	60.33	62.54	66.05	67.45	68.27	70.24	1.77	1.63	1.28	2.62	2.39	1.82
	E4	73.21	73.24	72.89	64.26	60.94	65.63	69.27	69.14	69.98	1.84	1.85	1.33	2.65	2.68	1.91
	E5	70.07	70.25	70.54	59.80	61.63	66.97	65.90	66.75	68.69	1.75	1.75	0.93	2.65	2.62	1.36
籽粒硬度 GH (%)	E1	75.02	69.37	75.17	51.99	49.51	55.72	61.66	58.76	65.97	4.99	4.36	4.09	8.10	7.42	6.20
	E2	76.16	72.29	73.20	53.60	51.13	53.75	66.02	59.51	67.24	5.14	5.36	4.09	7.79	9.00	6.07
	E3	69.44	67.07	70.07	53.35	49.54	54.14	61.56	56.27	64.42	3.92	3.97	3.34	6.37	7.05	5.18
	E4	77.10	70.26	68.89	53.03	49.70	52.78	68.67	57.63	63.44	5.38	4.57	3.57	7.83	7.92	5.62
	E5	70.30	73.12	70.65	45.98	48.64	53.29	57.97	55.55	63.51	5.56	4.28	3.66	9.59	7.70	5.76
出粉率 FY (%)	E1	74.36	73.44	75.22	62.39	57.47	66.46	68.42	69.57	72.06	2.71	2.19	1.82	3.96	3.14	2.52
	E2	80.24	77.06	76.73	68.37	68.37	71.28	73.09	72.46	74.78	2.00	1.55	1.27	2.73	2.14	1.70
	E3	77.41	74.34	74.16	67.10	66.82	69.34	71.12	69.89	72.51	1.59	1.41	1.22	2.24	2.02	1.68
	E4	76.55	75.40	76.93	68.18	67.33	69.93	72.43	70.84	73.18	1.77	1.55	1.34	2.44	2.19	1.82
	E5	75.38	74.65	73.72	67.38	64.45	67.53	71.20	68.55	70.51	1.68	1.56	1.34	2.36	2.28	1.90
容重 TW (g·L^-1)	E1	823.25	821.55	828.75	751.62	732.95	779.65	790.40	794.85	804.72	15.07	13.20	11.08	1.91	1.66	1.38
	E2	840.12	825.69	824.17	770.63	777.00	778.85	805.11	801.52	799.75	13.32	9.81	10.82	1.65	1.22	1.35
	E3	818.22	817.23	825.58	756.64	771.87	788.60	796.16	795.10	804.40	10.67	9.08	8.39	1.34	1.14	1.04
	E4	839.55	817.75	818.78	763.24	758.88	758.43	812.36	793.52	783.93	14.49	9.79	11.88	1.78	1.23	1.52
	E5	824.60	821.85	822.48	774.59	764.13	771.23	796.36	796.72	797.77	11.27	9.84	9.69	1.42	1.24	1.21

表1

表2

图1

图2

表3

春小麦品质性状显著关联的稳定的位点信息"

性状 Trait	位点 Marker	染色体 Chr.	物理位置 Position (Mb)	P值 P value	表型贡献率 R² (%)	环境 Environment
沉降值 SV (mL)	AX-109923777	1B	335.51	1.25E-05—4.48E-05	6.96—7.83	E4, E5
	AX-109389304	1B	445.67	6.44E-05—4.57E-04	5.49—6.79	E4, E5
	AX-94935157	1D	470.73—478.86	2.57E-05—3.85E-04	5.42—7.54	E2, E5
	AX-109486695	3A	725.15	1.15E-04—5.59E-04	5.11—6.23	E2, E5
	AX-108986146	3B	547.59	2.89E-05—3.18E-04	5.56—7.22	E4, E5
	AX-111583589	4A	121.51	5.34E-05—7.95E-04	4.95—6.94	E4, E5
	AX-110417316	4A	130.42	1.05E-05—6.22E-05	6.78—7.98	E4, E5
	AX-111264892	4A	241.73	1.40E-04—2.24E-04	5.79—6.17	E4, E5
	AX-109608730	4A	413.76	2.96E-05—3.05E-04	5.67—7.25	E4, E5
	AX-109285136	4A	594.90—598.79	9.75E-05—8.96E-04	3.82—6.39	E2, E3, E4
	AX-109834040	4A	609.82	2.7E-04—6.83E-04	4.98—5.65	E4, E5
	AX-110002719	4B	43.56	2.69E-04—9.30E-04	3.79—4.69	E2, E4
	AX-94513518	4B	556.06	2.13E-07—5.41E-04	5.24—10.89	E2, E3
	AX-110023236	5A	30.66	7.47E-04—9.28E-04	4.80—4.95	E2, E4
	AX-109374128	5A	600.17	7.45E-05—4.00E-04	5.43—6.61	E4, E5
	AX-110149809	5B	554.32	2.12E-04—3.56E-04	5.45—5.92	E3, E5
	AX-109983932	5B	604.61	5.60E-04—5.87E-04	5.15—5.18	E2, E3
	AX-109402563	5D	28.88	3.63E-05—1.86E-04	6.05—7.11	E4, E5
	AX-110082918	5D	529.22	5.79E-05—3.09E-04	5.57—6.72	E4, E5
	AX-109512342	6A	25.80—28.33	3.91E-05—4.89E-04	5.25—7.11	E2, E3, E4, E5
	AX-94564307	6D	21.98	4.64E-05—7.11E-04	4.98—6.93	E2, E4
	AX-111614875	6D	70.62	3.43E-05—1.81E-04	5.96—7.10	E4, E5
	AX-109452823	7A	26.47—32.70	5.93E-05—9.46E-04	3.72—6.34	E1, E2, E4, E5
	AX-109477546	7B	155.48—159.71	1.99E-05—7.71E-04	5.03—10.36	E2, E3, E4, E5
	AX-94420739	7D	540.87	9.04E-04—9.43E-04	4.83—5.08	E2, E4
出粉率 FY (%)	AX-109428041	1A	22.48	4.14E-04—7.46E-04	5.02—5.24	E3, E5
	AX-94802245	1D	73.73—77.36	1.14E-04—1.00E-03	4.88—6.34	E3, E4
	AX-111099555	2B	18.94—24.13	5.37E-05—3.64E-04	4.42—5.72	E3, E4
	AX-111040045	4A	632.76-659.66	1.51E-04—9.74E-04	3.83—5.96	E3, E5
	AX-110671478	6A	3.80—6.57	3.32E-06—9.73E-04	3.78—7.77	E3, E4
	AX-111534875	6A	5.51—12.28	1.44E-05—3.98E-04	5.61—7.91	E1, E3, E4
	AX-111745518	6A	17.54—24.41	1.51E-04—4.85E-04	6.14—6.15	E1, E4
	AX-111611885	6D	2.49—5.82	1.57E-04—7.67E-04	5.00—6.14	E3, E4
	AX-111076418	7A	8.91—12.44	1.70E-04—8.12E-04	3.75—5.14	E4, E5
蛋白质含量 PRC (%)	AX-94935157	1D	470.73—478.18	8.52E-04—9.52E-04	3.85—4.77	E2, E4
	AX-94694898	2B	620.64	2.57E-04—8.16E-04	4.87—5.71	E2, E4
	AX-111218282	2D	523.36	3.71E-04—8.94E-04	4.82—5.47	E2, E4
	AX-94589328	2D	624.10	6.02E-04—8.85E-04	5.10—5.12	E2, E4
	AX-110442313	3A	510.31—510.60	2.91E-04—8.05E-04	4.88—5.62	E2, E4
	AX-110002719	4B	43.56	1.20E-04—2.83E-04	4.59—5.13	E2, E4, E5
	AX-111560874	5B	10.50	1.38E-04—5.85E-04	5.11—6.15	E2, E4
	AX-109300733	6A	12.28—17.54	5.03E-05—8.33E-04	4.87—6.98	E2, E4
	AX-95194586	6D	77.52	3.12E-04—9.12E-04	4.97—5.68	E2, E4
	AX-111654743	7A	32.00—32.70	8.79E-05—8.48E-04	3.83—5.30	E1, E2, E3, E5
	AX-108797850	7B	453.77	2.79E-04—7.67E-04	4.93—5.66	E2, E4
	AX-108753131	7D	534.44—540.87	3.72E-04—6.37E-04	5.17—5.70	E2, E4
	AX-108974357	7D	566.20—566.23	4.25E-04—8.84E-04	4.99—5.36	E2, E4
	AX-110945986	7D	609.24—610.23	2.75E-04—8.28E-04	4.97—5.68	E2, E3
淀粉含量 STC (%)	AX-95180100	1D	359.67	2.12E-05—2.57E-04	5.75—7.55	E4, E5
	AX-94589328	2D	624.10—629.21	8.13E-05—8.67E-04	4.86—6.63	E1,E2
	AX-109934763	3D	559.16—561.01	3.24E-05—9.76E-04	4.78—7.56	E1, E2
	AX-110417316	4A	130.42—136.97	4.48E-04—8.12E-04	4.93—5.59	E1, E2
	AX-109332913	4A	701.53	2.32E-04—7.15E-04	5.26—5.79	E1, E2
	AX-110169414	5A	575.26	2.31E-04—5.72E-04	4.07—4.90	E1, E2
	AX-110046517	6A	8.09—12.28	2.58E-05—8.16E-04	5.14—7.43	E1, E3
	AX-111745518	6A	17.54	5.73E-04—8.12E-04	4.90—5.42	E1, E2
	AX-108888443	6D	16.23—21.98	6.92E-04—9.04E-04	4.82—5.26	E1, E2
	AX-110545157	7A	32.70	2.86E-04—7.03E-04	4.14—4.54	E1, E2
	AX-108974357	7D	566.20—566.23	2.60E-04—7.07E-04	5.00—5.80	E1, E2
容重 TW (g·L^-1)	AX-109301633	2A	84.15	4.89E-04—9.53E-04	4.50—5.44	E1, E3
	AX-110173580	2A	697.04—698.01	2.69E-04—4.71E-04	4.23—6.47	E1, E2
	AX-86184511	2B	158.15—164.43	2.03E-04—9.14E-04	4.53—4.79	E1, E4
	AX-110946610	4A	608.03—616.89	7.28E-04—7.84E-04	3.93—4.71	E1, E2
	AX-109947431	5B	672.48—678.51	2.29E-04—6.26E-04	5.26—5.84	E3, E5
	AX-110402876	7D	626.92	3.38E-04—4.10E-04	5.43—5.56	E2, E5
湿面筋含量 WCG (%)	AX-110002719	4B	43.56	1.27E-04—3.97E-04	4.32—5.06	E2, E4, E5
	AX-111560874	5B	10.50	2.83E-04—9.94E-04	4.71—5.61	E2, E4
	AX-111745518	6A	17.54—25.80	3.01E-04—6.74E-04	4.97—5.75	E2, E4
	AX-111654743	7A	32.00—32.70	7.64E-05—7.13E-04	4.72—5.35	E1, E2
	AX-94420739	7D	540.87	6.47E-04—6.52E-04	5.04—-5.22	E2, E4
籽粒硬度 GH (%)	AX-111107679	1A	476.08—483.94	1.32E-04—6.54E-04	4.92—5.01	E4, E5
	AX-110918237	1A	487.48—489.18	5.71E-05—8.41E-04	4.76—6.59	E4, E5
	AX-109499408	1A	496.31—500.66	1.77E-04—6.78E-04	3.94—5.91	E3, E5
	AX-110369038	1B	627.15—628.51	3.72E-04—9.45E-04	3.81—4.74	E1, E2, E3
	AX-94534666	1D	457.15—457.95	3.16E-06—5.85E-04	4.01—7.42	E2, E3, E5
	AX-111636440	2A	505.33—507.36	1.45E-04—9.12E-04	3.92—5.34	E1, E3
	AX-94447403	2D	439.44	5.38E-04—7.59E-04	4.84—5.39	E1, E3
	AX-111640432	3A	728.42—731.08	1.70E-04—5.00E-04	4.12—4.76	E2, E3, E5
	AX-110951756	3B	7.08	7.16E-06—8.58E-04	4.73—8.11	E2, E3
	AX-111018209	3D	553.59—560.75	3.97E-04—8.39E-04	4.98—5.28	E1, E5
	AX-111031457	5A	571.71—571.94	9.82E-05—1.82E-04	4.72—5.12	E3, E5
	AX-108945948	6A	9.14—13.06	1.00E-04—9.09E-04	5.00—6.02	E1, E5
	AX-108802871	7A	10.09—16.64	2.96E-04—8.09E-04	4.32—5.85	E1, E5
	AX-111664463	7A	123.20—127.55	8.90E-05—6.93E-04	3.86—6.29	E4, E5
	AX-110495496	7A	484.39—489.95	2.30E-04—9.39E-04	3.72—5.65	E3, E4, E5

表3

表4

同时与多个品质性状显著关联的稳定位点信息"

性状 Trait	位点 Marker	染色体 Chr.	物理位置 Position (Mb)	P值 P value	表型贡献率 R² (%)	环境 Environment
沉降值,蛋白质含量 SV, PRC	AX-94935157	1D	470.73—478.86	2.57E-05—9.52E-04	3.85—7.54	E2, E4, E5
淀粉含量,蛋白质含量 STC, PRC	AX-94589328	2D	624.10—629.21	8.13E-05—8.85E-04	4.86—6.63	E1, E2, E4
籽粒硬度,沉降值 GH, PRC	AX-111640432	3A	725.15—731.08	1.15E-04—5.59E-04	4.12—6.23	E2, E3, E5
籽粒硬度,淀粉含量 GH, STC	AX-111018209	3D	553.59—561.01	3.24E-05—9.76E-04	4.78—7.56	E1, E2, E5
淀粉含量,沉降值 STC, SV	AX-110417316	4A	130.42—136.97	1.05E-05—8.12E-04	4.93—7.98	E1, E2, E4, E5
容重,沉降值 TW, SV	AX-110946610	4A	608.03—616.89	2.70E-04—7.84E-04	3.93—5.65	E1, E2, E4, E5
沉降值,蛋白质含量,湿面筋含量 SV, PRC, WGC	AX-110002719	4B	43.56	1.20E-04—9.30E-04	3.79—5.13	E2, E4, E5
籽粒硬度,淀粉含量 GH, STC	AX-111031457	5A	571.71—575.26	9.82E-05—5.72E-04	4.07—5.12	E1, E2, E3, E5
蛋白质含量,湿面筋含量 PRC, WGC	AX-111560874	5B	10.50	1.38E-04—9.94E-04	4.71—6.15	E2, E4
出粉率,淀粉含量,籽粒硬度 FY, STC, GH	AX-111534875	6A	5.51—12.28	1.44E-05—9.09E-04	5.00—7.91	E1, E3, E4, E5
淀粉含量,蛋白质含量 STC, PRC	AX-111745518	6A	12.28—17.54	5.03E-05—8.33E-04	4.87—6.98	E1, E2, E4
出粉率,湿面筋含量 FY, WGC	AX-111745518	6A	17.54—25.80	1.51E-04—6.74E-04	4.97—6.15	E1, E2, E4
淀粉含量,沉降值 STC, SV	AX-108888443	6D	16.23—21.98	4.64E-05—9.04E-04	4.82—5.69	E1, E2, E4
沉降值,蛋白质含量 SV, PRC	AX-111614875	6D	70.62—77.52	3.43E-05—9.12E-04	4.97—7.10	E2, E4, E5
籽粒硬度,出粉率 GH, FY	AX-108802871	7A	8.91—16.64	1.70E-04—8.12E-04	3.75—5.85	E1, E4, E5
沉降值,淀粉含量,蛋白质含量,湿面筋含量 SV, STC, PRC, WGC	AX-109452823	7A	26.47—32.70	5.93E-05—9.46E-04	3.72—6.34	E1, E2, E3, E4, E5
沉降值,湿面筋含量,蛋白质含量 SV, WGC, PRC	AX-94420739	7D	534.44—540.87	3.72E-04—9.43E-04	4.83—5.70	E2, E4
蛋白质含量,淀粉含量 PRC, STC	AX-108974357	7D	566.20—566.23	2.60E-04—8.84E-04	4.99—5.80	E1, E2, E4

表4

表5

参考文献 40

[1]	王立祥, 廖允成. 中国粮食问题. 北京: 阳光出版社, 2012.
	WANG L X, LIAO Y C. Food Issues in China. Beijing: Sunshine Press, 2012. (in Chinese)
[2]	赵广才, 常旭虹, 王德梅, 陶志强, 王艳杰, 杨玉双, 朱英杰. 小麦生产概况及其发展. 作物杂志, 2018(4):1-7.
	ZHAO G C, CHANG X H, WANG D M, TAO Z Q, WANG Y J, YANG Y S, ZHU Y J. General situation and development of wheat production. Crops, 2018(4):1-7. (in Chinese)
[3]	燕丽, 王志忠, 郑文寅, 张文明, 郭文善, 姚大年. 基因型和环境对安徽小麦品质性状的影响. 麦类作物学报, 2016, 36(11):1497-1501.
	YAN L, WANG Z Z, ZHENG W Y, ZHANG W M, GUO W S, YAO D N. Effects of genotype and environment on wheat quality in Anhui province. Journal of Triticeae Crops, 2016, 36(11):1497-1501. (in Chinese)
[4]	关二旗, 魏益民, 张波, 郭进考, 张国权, 刘彦军, 罗勤贵, 班进福. 黄淮冬麦区部分区域小麦品种构成及品质性状分析. 中国农业科学, 2012, 45(6):1159-1168.
	GUAN E Q, WEI Y M, ZHANG B, GUO J K, ZHANG G Q, LIU Y J, LUO Q G, BAN J F. Analysis of the variety composition and quality properties of wheat in a part of the Yellow-Huai river zone. Scientia Agricultura Sinica, 2012, 45(6):1159-1168. (in Chinese)
[5]	张爱民, 李欣, 刘冬成, 孙家柱, 阳文龙. 品质支撑农作物产业与未来发展. 中国农业科学, 2016, 49(22):4265-4266.
	ZHANG A M, LI X, LIU D C, SUN J Z, YANG W L. Quality-the future of crop production. Scientia Agricultura Sinica, 2016, 49(22):4265-4266. (in Chinese)
[6]	郭利建, 王竹林, 汪世娟, 刘振华, 刘香利, 胡胜武, 赵惠贤. 基于SRAP和SSR标记的小麦品质相关性状的QTL定位. 麦类作物学报, 2016, 36(10):1275-1282.
	GUO L J, WANG Z L, WANG S J, LIU Z H, LIU X L, HU S W, ZHAO H X. QTL mapping of wheat grain quality traits based on SRAP and SSR marker. Journal of Triticeae Crops, 2016, 36(10):1275-1282. (in Chinese)
[7]	刘建军, 何中虎, 赵振东, 宋建民, 刘爱峰. 小麦面条加工品质研究进展. 麦类作物学报, 2001(2):81-84.
	LIU J J, HE Z H, ZHAO Z D, SONG J M, LIU A F. Review of noodle industrial quality of wheat. Journal of Triticeae Crops, 2001(2):81-84. (in Chinese)
[8]	陈锋, 何中虎, 崔党群, 赵武善, 张艳, 王德森. 利用近红外透射光谱技术测定小麦品质性状的研究. 麦类作物学报, 2003(3):1-4.
	CHEN F, HE Z H, CUI D Q, ZHAO W S, ZHANG Y, WANG D S. Measurement of wheat quality traits by near infrared transmittance spectroscopy. Journal of Triticeae Crops, 2003(3):1-4. (in Chinese)
[9]	关二旗, 魏益民, 张波. 小麦籽粒品质与基因型及环境条件的关系. 麦类作物学报, 2010, 30(5):963-969.
	GUAN E Q, WEI Y M, ZHANG B. Relationships between wheat kernel quality and genotype as well as environmental conditions. Journal of Triticeae Crops, 2010, 30(5):963-969. (in Chinese)
[10]	信志红, 郭建平, 谭凯炎, 刘凯文, 杨荣光, 张利华, 孙义. 冬小麦籽粒品质评价及其对气象因子的响应研究. 中国生态农业学报, 2019, 27(8):1205-1217.
	XIN Z H, GUO J P, TAN K Y, LIU K W, YANG R G, ZHANG L H, SUN Y. Evaluation of grain quality of winter wheat and its response to meteorological factors. Chinese Journal of Eco-Agriculture, 2019, 27(8):1205-1217. (in Chinese)
[11]	NELSON J C, ANDREESCU C, BRESEGHELLO F, FINNEY P L, GUALBERTO D G, BERGMAN C, PENA R J, PERRETANT M R, LEROY P, QUALSET C O, SORRELLS M E. Quantitative trait locus analysis of wheat quality traits. Euphytica, 2006, 149:145-159. doi: 10.1007/s10681-005-9062-7
[12]	TURUSPEKOV Y, BAIBULATOVA A, YENMEKBAYEY K, TOKHETOVA L, CHUDINOV V, SEREDA G, GANAL M, GRIFFITHS S, ABUGALIEVA S. GWAS for plant growth stages and yield components in spring wheat (Triticum aestivum L.) harvested in three regions of Kazakhstan. BMC Plant Biology, 2017, 17(S1):51-61. doi: 10.1186/s12870-017-0999-1
[13]	LI H M, TANG Z X, ZHANG H Q, YAN B J, REN Z L. Major quality trait analysis and QTL detection in hexaploid wheat in humid rain-fed agriculture. Genetics and Molecular Research, 2013, 12(2):1740-1751. doi: 10.4238/2013.May.21.5
[14]	ECHEVENY-SOLARTE M, KUMAR A, KIANIAN S, SIMSEK S, ALAMRI M S, MANTOVANI E E, MCCLEAN P E, DECKARD E L, ELIAS E, SCHATZ B, XU S S, MERGOUM M. New QTL alleles for quality-related traits in spring wheat revealed by RIL population derived from supernumerary×non-supernumerary spikelet genotypes. Theoretical and Applied Genetics, 2015, 128(5):893-912. doi: 10.1007/s00122-015-2478-0
[15]	吴云鹏, 张业伦, 肖永贵, 阎俊, 张勇, 张晓科, 张利民, 夏先春, 何中虎. 小麦重要品质性状的QTL定位. 中国农业科学, 2008, 41(2):331-339.
	WU Y P, ZHANG Y L, XIAO Y G, YAN J, ZHANG Y, ZHANG X K, ZHANG L M, XIA X C, HE Z H. QTL mapping for important quality traits in common wheat. Scientia Agricultura Sinica, 2008, 41(2):331-339. (in Chinese)
[16]	黄梦豪, 刘天相, 强琴琴, 李春莲, 王中华. 基于SNP和SSR标记的小麦品质性状的QTL定位. 分子植物育种, 2019, 17(12):3966-3973.
	HUANG M H, LIU T X, QIANG Q Q, LI C L, WANG Z H. QTL mapping of wheat grain quality traits based on SNP and SSR marker. Molecular Plant Breeding, 2019, 17(12):3966-3973. (in Chinese)
[17]	WURSCHUM T, LANGER S M, LONGIN C F, KORZUN V, AKHUNOV E, EBMEYER E, SCHACHSCHNEIDER R, SCHACHT J, KAZMAN E, REIF J C. Population structure, genetic diversity and linkage disequilibrium in elite winter wheat assessed with SNP and SSR markers. Theoretical and Applied Genetics, 2013, 126(6):1477-1486. doi: 10.1007/s00122-013-2065-1
[18]	MENG L, LI H, ZHANG L, WANG J. QTL IciMapping: Integrated software for genetic linkage map construction and quantitative trait locus mapping in biparental populations. The Crop Journal, 2015, 3(3):269-283. doi: 10.1016/j.cj.2015.01.001
[19]	陈昆松, 李方, 徐昌杰, 张上隆, 傅承新. 改良CTAB法用于多年生植物组织基因组DNA的大量提取. 遗传, 2004, 26(4):529-531.
	CHEN K S, LI F, XU C J, ZHANG S L, FU C X. Modified CTAB method for extraction of genome DNA from perennial plant tissues. Hereditas, 2004, 26(4):529-531. (in Chinese)
[20]	WANG S X, ZHU Y L, ZHANG D X, SHAO H, LIU P, HU J B, ZHANG H, ZHANG H P, CHANG C, LU J, XIA X C, SUN Z L, MA C X. Genome-wide association study for grain yield and related traits in elite wheat varieties and advanced lines using SNP markers. PLoS ONE, 2017, 12(11):1-14.
[21]	ZHU C S, GORE M, BUCKLER E S, BUCKLER, YU J M. Status and prospects of association mapping in plants. The Plant Genome, 2008, 1(1):5-20.
[22]	YU J, PRESSOIR G, BRIGGS W H, BI I V, YAMASAKI M, DOEBIEY J F, MCMULLEN M D, GAUT B S, NIELSEN D M, HOLLAND J B, KRESOVICH S, BUCKLER E S. A unified mixed-model method for association mapping that accounts for multiple levels of relatedness. Nature Genetics, 2006, 38(2):203-208. doi: 10.1038/ng1702
[23]	翟俊鹏, 李海霞, 毕惠惠, 周思远, 罗肖艳, 陈树林, 程西永, 许海霞. 普通小麦主要农艺性状的全基因组关联分析. 作物学报, 2019, 45(10):1488-1502.
	ZHAI J P, LI H X, BI H H, ZHOU S Y, LUO X Y, CHEN S L, CHENG X Y, XU H X. Genome-wide association study for main agronomic traits in common wheat. Acta Agronomica Sinica, 2019, 45(10):1488-1502. (in Chinese)
[24]	BRESEGHELLO F, SORRELLS M E. Association mapping of kernel size and milling quality in wheat (Triticum aestivum L.) cultivars. Genetics, 2006, 172(2):1165-1177. doi: 10.1534/genetics.105.044586
[25]	周思远, 毕惠惠, 程西永, 张旭睿, 闰永行, 王航辉, 毛培钧, 李海霞, 许海霞. 小麦耐低磷相关性状的全基因组关联分析. 植物遗传资源学报, 2020, 21(2):431-445.
	ZHOU S Y, BI H H, CHENG X Y, ZHANG X R, RUN Y H, WANG H H, MAO P J, LI H X, XU H X. Genome-wide association study of low-phosphorus tolerance related traits in wheat. Journal of Plant Genetic Resources, 2020, 21(2):431-445. (in Chinese)
[26]	张金波, 严勇亮, 王小波, 路子峰, 肖菁, 彭惠茹, 丛花. 新疆春小麦育成品种遗传演变分析. 新疆农业科学, 2020, 57(3):418-426.
	ZHANG J B, YAN Y L, WANG X B, LU Z F, XIAO J, PENG H R, CONG H. Analysis of the genetic evolution of cultivated spring wheat varieties in Xinjiang. Xinjiang Agricultural Sciences, 2020, 57(3):418-426. (in Chinese)
[27]	金欣欣, 姚艳荣, 贾秀领, 姚海坡, 申海平, 崔永增, 李谦. 基因型和环境对小麦产量、品质和氮素效率的影响. 作物学报, 2019, 45(4):635-644.
	JIN X X, YAO Y R, JIA X R, YAO H B, SHEN H P, CUI Y Z, LI Q. Effects of genotype and environment on wheat yield, quality, and nitrogen use efficiency. Acta Agronomica Sinica, 2019, 45(4):635-644. (in Chinese)
[28]	张影全, 唐娜, 张波, 景东林, 马永安, 魏益民. 冀南地区小麦籽粒品质现状及利用潜力分析. 麦类作物学报, 2018, 38(2):157-163.
	ZHANG Y Q, TANG N, ZHANG B, JING D L, MA Y A, WEI Y M. Study on kernel quality property and processing potency of wheat in the south of Hebei province. Journal of Triticeae Crops, 2018, 38(2):157-163. (in Chinese)
[29]	吴新元, 芦静, 张新忠, 黄天荣, 李建疆, 周安定, 梁晓东, 曹俊梅, 高永红, 曾潮武. 新疆小麦品质生态区划研究. 新疆农业科学, 2017, 54(8):1373-1383.
	WU X Y, LU J, ZHANG X Z, HUANG T R, LI J J, ZHOU A D, LIANG X D, CAO J M, GAO Y H, ZENG C W. Study of ecological division for wheat quality in Xinjiang. Xinjiang Agricultural Sciences, 2017, 54(8):1373-1383. (in Chinese)
[30]	杨林, 吴青霞, 邵慧, 冉从福, 余静, 李立群, 李学军. 小麦籽粒品质性状的QTL分析. 西北植物学报, 2013, 33(8):1574-1583.
	YANG L, WU Q X, SHAO H, RAN C F, YU J, LI L Q, LI X J. QTL mapping for grain quality traits in wheat. Acta Botanica Boreali-Occidentalia Sinica, 2013, 33(8):1574-1583. (in Chinese)
[31]	LOU H Y, ZHANG R Q, LIU Y T, GUO D D, ZHAI S S, CHEN A Y, ZHANG Y F, XIE C J, YOU M S, PENG H R, LIANG R Q, NI Z F, SUN Q X, LI B Y. Genome-wide association study of six quality-related traits in common wheat (Triticum aestivum L.) under two sowing condition. Theoretical and Applied Genetics, 2021, 134:399-418 doi: 10.1007/s00122-020-03704-y
[32]	沈玮囡, 王竹林, 杨睿, 李美霞, 梁子英, 奚亚军, 孙风丽, 刘曙东. 波兰小麦品系XN555×普通小麦品系中13衍生重组自交系(RILs)群体中籽粒品质相关性状QTL定位. 农业生物技术学报, 2014, 22(5):561-571.
	SHEN W N, WANG Z L, YANG R, LI M X, LIANG Z Y, XI Y J, SUN F L, LIU S D. QTL analysis of grain quality related traits using recombinant inbred lines (RILs) derived from the cross of Triticum polonicum L. line XN555×T. aestivum L. line Zhong 13. Journal of Agricultural Biotechnology, 2014, 22(5):561-571. (in Chinese)
[33]	贾鑫磊, 何贝轩, 郭丹丹, 郭美丽. 膨胀素和木葡聚糖内转葡糖基酶/水解酶基因的功能研究进展. 植物生理学报, 2018, 54(11):1659-1668.
	JIA X L, HE B X, GUO D D, GUO M L. Research progress in the function of expansins and xyloglucan endotransglucosylase/hydrolase. Plant Physiology Journal, 2018, 54(11):1659-1668. (in Chinese)
[34]	MIEDES E, ZARRA I, HOSON T, HERBERS K, SONNEWALD U, LORENCES E P. Xyloglucan endotransglucosylase and cell wall extensibility. Journal of Plant Physiology, 2011, 168(3):196-203. doi: 10.1016/j.jplph.2010.06.029
[35]	XU P, CAI X T, WANG Y, XING L, CHEN Q, XIANG C B. HDG11 upregulates cell-wall-loosening protein genes to promote root elongation in Arabidopsis. Journal of Experimental Botany, 2014, 65(15):4285-4295. doi: 10.1093/jxb/eru202
[36]	HAN Y, HAN S, BAN Q, HE Y H, JIN M J, RAO J P. Overexpression of persimmon DkXTH1 enhanced tolerance to abiotic stress and delayed fruit softening in transgenic plants. Plant Cell Reports, 2017, 36:583-596. doi: 10.1007/s00299-017-2105-4
[37]	茹京娜, 于太飞, 陈隽, 陈明, 周永斌, 马有志, 徐兆师, 闵东红. 小麦锌指转录因子TaDi19A对低温的响应及其互作蛋白的筛选. 中国农业科学, 2017, 50(13):2411-2422.
	RU J N, YU T F, CHEN Y, CHEN M, ZHOU Y B, MA Y Z, XU Z S, MIN D H. TaDi19A of zinc finger transcription factors in wheat in response to low temperature. Scientia Agricultura Sinica, 2017, 50(13):2411-2422. (in Chinese)
[38]	魏春茹, 孟钰玉, 范润侨, 赵梦伊, 于秀梅, 赵伟全, 康振生, 刘大群. 小麦F-box/Kelch类基因TaFKOR23的抗逆相关表达模式及分子互作蛋白鉴定. 植物遗传资源学报, 2020, 21(3):695-705.
	WEI C R, MENG Y Y, FAN R Q, ZHAO M Y, YU X M, ZHAO W Q, KANG Z S, LIU D Q. Stress-related expression profile of F-box/Kelch gene TaFKOR23 in wheat and molecular characterization of the interacting target protein. Journal of Plant Genetic Resources, 2020, 21(3):695-705. (in Chinese)
[39]	邱丽丽, 赵琪, 张玉红, 戴绍军. 植物质膜蛋白质组的逆境应答研究进展. 植物学报, 2017, 52(2):128-147. doi: 10.11983/CBB16001
	QIU L L, ZHAO Q, ZHANG Y H, DAI S J. Advances in adversity response in proteome of plant plasma membrane. Chinese Bulletin of Botany, 2017, 52(2):128-147. (in Chinese) doi: 10.11983/CBB16001
[40]	陈雪燕, 王灿国, 程敦公, 李豪圣, 宋健民, 刘爱峰, 王利彬, 董爽爽, 赵振东, 刘建军, 曹新有. 小麦加工品质相关贮藏蛋白、基因及其遗传改良研究进展. 植物遗传资源学报, 2018, 19(1):1-9.
	CHEN X Y, WANG C G, CHENG D G, LI H S, SONG J M, LIU A F, WANG L B, DONG S S, ZHAO Z D, LIU J J, CAO X Y. Research progress on wheat processing quality related storage proteins, genes and genetic improvement. Journal of Plant Genetic Resources, 2018, 19(1):1-9. (in Chinese)

性状 Trait	基因型 Genotype	环境 Environment	基因型×环境 G×E	重复 Replicate	误差 Error	遗传力 H²
蛋白质含量PRC	30.50***	323.25***	2.71***	28.92***	0.94***	0.92
湿面筋含量WGC	173.71***	1575.75***	14.49***	138.56***	5.07***	0.92
沉降值SV	396.29***	5503.76***	38.53***	482.08***	12.76***	0.91
淀粉含量STC	42.33***	1067.05***	4.58***	50.85***	1.38***	0.90
籽粒硬度GH	406.19***	4375.56***	20.43***	20.52***	3.83***	0.95
出粉率FY	41.49***	1793.73***	4.76***	5.17***	0.91***	0.89
容重TW	1308.20***	7599.86***	267.46***	247.03***	54.04***	0.82