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.

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

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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