陕A群、陕B群选育玉米自交系及其杂交种种子活力评价及遗传解析

doi:10.3864/j.issn.0578-1752.2026.16.002

摘要/Abstract

摘要：

【目的】玉米种子活力是决定种子品质的关键因素之一，直接影响出苗率、幼苗生长及群体整齐度，进而影响产量形成。筛选高活力自交系与杂交种，挖掘调控种子萌发的关键基因，为培育高活力玉米品种提供理论依据。【方法】选取陕A群、陕B群2个杂种优势群选育的22份玉米自交系并采用NC-Ⅱ遗传设计组配104份F₁杂交组合。通过标准发芽试验和冷浸发芽试验测定发芽势、发芽率、发芽指数、苗长、根长、苗干重，以及活力指数Ⅰ、Ⅱ、Ⅲ共9个性状，开展种子活力的综合评价。结合基因型数据与表型数据，对主要性状开展全基因组关联分析，筛选与种子活力显著相关的数量性状核苷酸位点，并整合公共转录组数据对潜在候选基因进行挖掘与功能预测。【结果】自交系与杂交种的种子活力相关性状均存在显著遗传变异。相较于标准发芽，冷浸处理显著降低了发芽势、发芽率和发芽指数，同时扩大了表型变异幅度，增强了基因型间的差异表达。多数性状在基因型、处理及其互作效应上达到显著或极显著水平，证实种子活力受遗传因素显著调控。发芽性状与苗期生长性状和活力指数呈显著正相关。并筛选出一批在不同发芽条件下表现稳定、对冷浸处理敏感性较低的优良材料，其中，在冷浸条件下，以陕A群的KA105、KA088、KA085、KA205和陕B群的KB088、KB060、KB168、KB076、KB021为亲本组配的杂交组合仍保持较高种子活力水平。自身活力中等的自交系在特定组合中可显著提升后代表现。全基因组关联分析共检测到43个与种子活力显著相关的QTN位点，其中7个为冷浸条件下的多性状共定位位点。利用公共转录组数据进一步筛选出11个候选基因，其中8个具有功能注释，主要涉及胁迫响应与转录调控等生物学过程。【结论】玉米种子活力为遗传控制的数量性状，基因型变异是其表型差异的主要来源。种子活力受多位点协同调控，低温条件下存在关键遗传区域参与调节。有利等位基因的积累有助于提升杂交种活力，同时杂交种表现还受亲本间特异遗传互作影响。

关键词: 玉米, 种子活力, 自交系, 杂交种, 全基因组关联分析

Abstract:

【Objective】 Seed vigor is one of the key determinants of seed quality in maize, directly influencing emergence rate, seedling growth, and stand uniformity, thereby affecting final yield formation. This study aimed to identify high-vigor inbred lines and hybrids, uncover key genes involved in regulating seed germination, and provide a theoretical basis for breeding high-vigor maize varieties.【Method】 Twenty-two maize inbred lines from two heterotic groups, Shaan A Group and Shaan B Group, were used to produce 104 F₁ hybrids following an NC-Ⅱ mating design. Seed vigor was evaluated under both the standard germination test and the cold soaking germination test. Nine traits were recorded, including germination energy (GE), germination percentage (GP), germination index (GI), seedling length (SL), root length (RL), dry weight (DW), vigor index I (VIⅠ), vigor index Ⅱ (VIⅡ), and vigor index Ⅲ (VIⅢ). Genotypic and phenotypic data were combined to perform a genome-wide association study (GWAS) for major traits. Quantitative trait nucleotides (QTNs) significantly associated with seed vigor were identified. Public transcriptome data were further used to screen potential candidate genes and to predict their possible functions. 【Result】 Significant genetic variation was observed for seed vigor-related traits in both inbred lines and hybrids. Compared with the standard germination test, cold soaking treatment reduced GE, GP, and GI, and increased the range of phenotypic variation among genotypes. Most traits showed significant effects of genotype, treatment, and their interaction, indicating that seed vigor is mainly controlled by genetic factors. Germination traits were positively correlated with seedling growth traits and vigor indices. Several materials showed stable performance under different germination conditions and low sensitivity to cold soaking. Hybrids derived from KA105, KA088, KA085, and KA205 of the Shaan A Group, and KB088, KB060, KB168, KB076, and KB021 of the Shaan B Group maintained relatively high seed vigor under cold soaking conditions. Some inbred lines with moderate seed vigor were able to improve hybrid performance in specific combinations. GWAS detected 43 QTNs significantly associated with seed vigor, including seven loci that were shared by two or more traits under cold soaking conditions. Based on transcriptome data, 11 candidate genes were identified, of which eight had functional annotation and were mainly related to stress response and transcription regulation.【Conclusion】 Maize seed vigor is a quantitative trait under genetic control, and genetic variation is the main source of phenotypic differences. Seed vigor is regulated by multiple loci, with key genomic regions involved under low-temperature conditions. The accumulation of favorable alleles can improve seed vigor in hybrids, and hybrid performance is also influenced by specific genetic interactions between parents.

Key words: maize, seed vigor, inbred line, hybrids, genome-wide association analysis

曹凡, 唐瑶, 王孟玮, 张栋渊, 卫亚博, 吴爽, 朱万超, 徐淑兔, 张兴华, 薛吉全. 陕A群、陕B群选育玉米自交系及其杂交种种子活力评价及遗传解析[J]. 中国农业科学, 2026, 59(16): 3476-3495.

CAO Fan, TANG Yao, WANG MengWei, ZHANG DongYuan, WEI YaBo, WU Shuang, ZHU WanChao, XU ShuTu, ZHANG XingHua, XUE JiQuan. Evaluation and Genetic Analysis of Seed Vigor in Inbred Lines and Responding Hybrids in Shaan A and Shaan B Groups[J]. Scientia Agricultura Sinica, 2026, 59(16): 3476-3495.

图/表 15

表1

表2

自交系与杂交种种子活力相关性状的描述性统计"

材料 Material	性状 Trait	处理 Treatment	变异范围 Range	平均值±标准偏差 Mean±SD	偏度 Skewness	峰度 Kurtosis	变异系数 CV (%)
自交系 Inbred lines	发芽势GE (%)	SGT	53.33-100.00	85.35±0.13	-0.99	0.32	15.34
	发芽势GE (%)	CST	4.44-95.56	58.64±0.24	-0.37	-0.48	41.50
	发芽率GP (%)	SGT	84.44-100.00	96.26±0.05	-1.27	0.51	5.02
	发芽率GP (%)	CST	8.89-98.89	65.40±0.24	-0.57	-0.23	36.47
	发芽指数GI	SGT	6.14-9.81	7.74±0.92	0.48	-0.19	11.91
	发芽指数GI	CST	0.60-9.11	5.56±2.22	-0.37	-0.45	39.93
	苗长SL (cm)	SGT	6.79-12.59	9.57±1.62	-0.04	-0.83	16.88
	苗长SL (cm)	CST	3.88-12.74	9.99±2.05	-1.13	2.31	20.52
	根长RL (cm)	SGT	14.54-21.34	17.07±1.50	0.93	1.97	8.79
	根长RL (cm)	CST	4.28-22.49	16.90±3.68	-1.78	6.08	21.75
	苗干重DW (g)	SGT	0.14-0.74	0.51±0.13	-0.94	1.99	25.59
	苗干重DW (g)	CST	0.46-0.88	0.68±0.12	-0.10	-0.74	17.18
	活力指数I VIⅠ	SGT	3.08-8.15	5.37±1.41	0.43	-0.47	26.32
	活力指数I VIⅠ	CST	0.08-5.53	3.03±1.50	0.04	-1.17	51.97
	活力指数Ⅱ VIⅡ	SGT	49.16-114.95	75.09±19.86	0.38	-0.91	26.44
	活力指数Ⅱ VIⅡ	CST	2.33-99.17	58.63±28.29	-0.22	-1.07	48.25
	活力指数Ⅲ VI Ⅲ	SGT	98.21-209.28	132.93±25.12	1.32	3.01	18.90
	活力指数Ⅲ VI Ⅲ	CST	2.57-204.75	99.59±49.24	0.11	-0.36	49.44
杂交种 Hybrids	发芽势GE (%)	SGT	55.56-100.00	90.55±0.10	-1.67	2.26	11.49
	发芽势GE (%)	CST	31.11-97.78	74.79±0.15	-0.95	0.31	19.88
	发芽率GP (%)	SGT	85.56-100.00	98.17±0.03	-2.47	6.69	2.85
	发芽率GP (%)	CST	36.67-100.00	80.79±0.13	-1.04	0.85	16.70
	发芽指数GI	SGT	6.29-9.94	8.25±0.94	-0.01	-1.10	11.40
	发芽指数GI	CST	3.18-9.64	7.21±1.34	-0.82	0.25	18.64
	苗长SL (cm)	SGT	8.47-16.69	12.61±1.84	-0.14	-0.68	14.62
	苗长SL (cm)	CST	4.95-15.97	11.89±1.87	-0.80	1.18	15.72
	根长RL (cm)	SGT	15.45-25.68	19.71±1.88	0.46	0.27	9.53
	根长RL (cm)	CST	11.53-22.40	18.18±1.82	-0.62	1.13	10.01
	苗干重DW (g)	SGT	0.46-1.17	0.86±0.13	-0.29	0.43	15.36
	苗干重DW (g)	CST	0.37-1.06	0.71±0.13	0.25	0.33	17.78
	活力指数ⅠVIⅠ	SGT	3.38-11.08	7.12±1.56	0.05	-0.27	21.89
	活力指数ⅠVIⅠ	CST	1.56-8.25	5.20±1.49	-0.30	-0.30	28.65
	活力指数Ⅱ VIⅡ	SGT	53.33-153.60	105.20±24.94	0.20	-0.96	23.71
	活力指数Ⅱ VIⅡ	CST	20.98-143.65	87.06±24.67	-0.41	-0.21	28.34
	活力指数Ⅲ VI Ⅲ	SGT	100.33-228.19	163.61±30.51	0.31	-0.81	18.65
	活力指数Ⅲ VI Ⅲ	CST	48.85-210.06	132.60±33.00	-0.44	0.07	24.89

表2

图1

图2

表3

图3

图4

表4

图5

图6

图7

表5

全基因组关联分析检测到的重要关联位点信息"

序号 No.	标记 Marker	染色体 Chromosome	位置 Position (bp)	基因型 Genotype	模型 Model	性状 Trait	P值 P value	表型解释率 R²(%)
SNP1	1_3353289	1	3353289	G/A	Add	SGT-GI	9.00E-06	11.43
SNP2	1_43499054	1	43499054	T/C	Rec	CST-GP	1.24E-10	18.26
SNP3	1_84304371	1	84304371	C/T	Rec	CST-SL	2.86E-12	25.07
SNP4	1_85839980	1	85839980	A/C	Rec	SGT-SL	2.05E-21	32.68
SNP5	1_154295033	1	154295033	A/G	Add	CST-GE	7.82E-08	17.78
SNP6	1_178647142	1	178647142	A/G	Rec	SGT-DW	1.55E-12	22.04
SNP7	1_291856974	1	291856974	T/C	Dom	SGT-SL	5.13E-15	22.47
SNP8	2_18197441*	2	18197441	A/G	Dom	CST-GE	6.97E-15	18.78
					Dom	CST-GI	2.80E-15	19.43
					Dom	CST-GP	1.10E-19	26.68
					Add	CST-GI	9.57E-15	18.69
SNP9	2_18739525	2	18739525	T/C	Dom	SGT-DW	2.63E-18	13.73
SNP10	2_19244047	2	19244047	T/C	Rec	SGT-GI	3.47E-09	17.49
SNP11	2_19372692	2	19372692	C/T	Add	CST-GE	8.77E-16	21.42
SNP12	2_30802598	2	30802598	G/C	Add	SGT-GE	6.48E-08	19.44
SNP13	2_39940294	2	39940294	G/A	Dom	SGT-RL	1.25E-06	11.55
SNP14	2_183378697	2	183378697	C/T	Dom	SGT-GI	6.59E-10	13.51
SNP15	3_17590250*	3	17590250	G/A	Dom	CST-RL	6.46E-10	17.11
					Dom	CST-SL	3.85E-15	24.91
SNP16	3_17805407*	3	17805407	C/T	Rec	CST-GE	5.57E-18	35.60
					Rec	CST-GI	8.49E-18	30.14
					Rec	CST-GP	7.32E-16	24.59
SNP17	3_17932972*	3	17932972	T/C	Dom	CST-GE	4.91E-16	24.05
					Dom	CST-GI	1.28E-14	20.89
SNP18	3_19091776	3	19091776	G/A	Dom	CST-GP	8.60E-11	11.02
SNP19	3_22825936	3	22825936	C/G	Dom	SGT-SL	1.34E-14	23.70
SNP20	3_32242592	3	32242592	C/A	Dom	SGT-DW	1.50E-07	12.62
SNP21	3_155021348	3	155021348	A/C	Dom	SGT-GE	3.33E-27	19.80
SNP22	3_192210299	3	192210299	G/A	Rec	SGT-RL	1.04E-06	19.69
SNP23	4_15716701	4	15716701	C/A	Rec	CST-DW	3.26E-05	12.32
SNP24	4_18511010*	4	18511010	G/C	Rec	CST-GE	6.81E-07	13.60
					Rec	CST-GI	1.31E-08	15.92
SNP25	4_28100027	4	28100027	C/A	Dom	CST-DW	3.08E-12	12.23
SNP26	4_52506095	4	52506095	C/T	Add	CST-SL	3.61E-10	32.65
SNP27	4_69862542	4	69862542	C/T	Add	SGT-GI	2.17E-06	16.18
SNP28	4_145757636	4	145757636	G/C	Dom	SGT-GE	8.07E-16	18.45
SNP29	5_7270274	5	7270274	G/A	Rec	SGT-GP	6.93E-06	15.31
SNP30	5_11611314	5	11611314	C/A	Add	SGT-SL	3.07E-18	47.37
SNP31	5_11960070	5	11960070	T/C	Rec	SGT-DW	2.63E-10	17.35
SNP32	5_142175693	5	142175693	A/G	Dom	CST-DW	7.30E-18	14.83
SNP33	5_204262820*	5	204262820	T/A	Rec	CST-GI	3.49E-06	8.56
					Rec	CST-GP	2.44E-09	14.74
					Add	CST-GI	5.57E-13	14.79
					Add	CST-GP	5.80E-08	14.18
					Add	CST-GE	1.63E-08	10.12
SNP34	6_96096697	6	96096697	C/T	Rec	CST-SL	3.58E-07	13.13
SNP35	6_159316307	6	159316307	C/T	Add	SGT-GE	6.71E-07	11.82
SNP36	6_168385013	6	168385013	C/T	Dom	SGT-GI	1.57E-08	10.40
SNP37	7_21288129	7	21288129	G/A	Dom	CST-DW	2.01E-18	13.75
SNP38	8_170405872	8	170405872	C/A	Add	SGT-RL	5.22E-08	22.64
SNP39	9_149806469	9	149806469	C/A	Add	CST-DW	1.12E-09	15.24
SNP40	9_153233111	9	153233111	C/T	Dom	CST-SL	3.84E-12	18.95
SNP41	10_9606210	10	9606210	T/C	Dom	SGT-GI	1.93E-07	11.63
SNP42	10_111383922	10	111383922	A/C	Rec	SGT-GI	5.59E-07	10.33
SNP43	10_138859255*	10	138859255	C/T	Dom	CST-GE	6.04E-08	7.74
					Dom	CST-GI	7.85E-10	10.36
					Dom	CST-GP	1.35E-07	6.76

表5

图8

图9

表6

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序号No.	材料名称Name	来源Source	序号No.	材料名称Name	来源Source
M1	KA105	陕A群Shaan A Group	P4	KB060	陕B群Shaan B Group
M2	KA023	陕A群Shaan A Group	P5	KB076	陕B群Shaan B Group
M3	KA205	陕A群Shaan A Group	P6	KB077-2	陕B群Shaan B Group
M4	KA215	陕A群Shaan A Group	P7	KB088	陕B群Shaan B Group
M5	KA085	陕A群Shaan A Group	P8	KB017	陕B群Shaan B Group
M6	KA088	陕A群Shaan A Group	P9	KB113	陕B群Shaan B Group
M7	KB-9-1	陕B群Shaan B Group	P10	KB168	陕B群Shaan B Group
M8	KB-9-2	陕B群Shaan B Group	P11	KB177	陕B群Shaan B Group
P1	KB017	陕B群Shaan B Group	P12	KB179	陕B群Shaan B Group
P2	KB021	陕B群Shaan B Group	P13	KB209	陕B群Shaan B Group
P3	KB042-2	陕B群Shaan B Group	P14	KB562	陕B群Shaan B Group

材料 Material	性状 Trait	基因型 Genotype			处理 Treatment			处理×基因型 Genotype×Treatment
材料 Material	性状 Trait	自由度 DF	均方 MS	F值 F-value	自由度 DF	均方 MS	F值 F-value	自由度 DF	均方 MS	F值 F-value
自交系 Inbred lines	发芽势GE	21	0.14	36.83**	1	2.36	605.78**	21	0.09	22.08**
	发芽率GP	21	0.11	39.73**	1	3.14	1168.19**	21	0.07	26.38**
	发芽指数GI	21	10.64	48.18**	1	159.99	724.255**	21	6.57	29.73**
	苗长SL	21	15.33	14.83**	1	5.37	5.19	21	5.30	5.13
	根长RL	21	33.27	32.05**	1	0.65	0.63	21	11.67	11.24**
	苗干重DW	21	0.08	31.35**	1	0.99	384.99**	21	0.01	4.57
	活力指数Ⅰ VIⅠ	21	11.01	27.52**	1	75.22	188.09**	21	4.39	10.97**
	活力指数Ⅱ VIⅡ	21	2409.15	28.49**	1	9115.14	107.78**	21	1124.84	13.30**
	活力指数Ⅲ VI Ⅲ	21	6158.26	43.45**	1	37043.47	263.76**	21	2900.96	20.66**
杂交种 Hybrids	发芽势GE	105	0.05	16.72**	1	3.95	1326.67**	105	0.05	16.49**
	发芽率GP	105	0.03	19.51**	1	4.80	2917.90**	105	0.02	15.11**
	发芽指数GI	105	4.60	18.73**	1	175.54	713.92**	105	3.67	14.93**
	苗长SL	105	17.50	13.93**	1	83.48	66.48	105	3.18	2.54**
	根长RL	105	13.36	7.08**	1	374.04	198.29**	105	7.17	3.80*
	苗干重DW	105	0.08	16.23**	1	3.35	676.20**	105	0.02	4.04**
	活力指数ⅠVIⅠ	105	8.72	15.24**	1	252.55	441.49**	105	4.35	7.61**
	活力指数Ⅱ VIⅡ	105	2687.82	18.32**	1	53441.35	364.15**	105	1038.37	7.08**
	活力指数Ⅲ VI Ⅲ	105	3894.52	15.62**	1	154790.67	620.68**	105	2248.23	9.02**

材料 Materials	类别 Type	材料数量 Number	发芽势 GE (%)		发芽率 GP (%)		发芽指数 GI		活力指数Ⅰ VIⅠ		活力指数Ⅱ VIⅡ		活力指数Ⅲ VIⅢ
材料 Materials	类别 Type	材料数量 Number	SGT	CST	SGT	CST	SGT	CST	SGT	CST	SGT	CST	SGT	CST
自交系 Inbred lines	第Ⅰ类Category Ⅰ	8	88.33	83.61	99.31	89.03	7.91	7.87	5.80	4.79	78.32	87.97	136.13	150.27
	第Ⅱ类Category Ⅱ	9	82.34	55.06	94.44	63.21	7.63	5.21	5.02	2.53	73.63	53.39	131.43	89.17
	第Ⅲ类Category Ⅲ	5	86.00	25.10	94.67	31.95	7.69	2.46	5.27	1.10	72.53	21.14	130.51	37.26
杂交种 Hybrids	第Ⅰ类Category Ⅰ	51	83.70	84.51	98.21	89.69	7.86	8.06	6.68	5.98	97.26	99.24	150.65	152.22
	第Ⅱ类Category Ⅱ	41	93.28	71.65	98.51	77.75	8.64	6.96	7.66	4.97	114.94	84.33	177.48	125.66
	第Ⅲ类Category Ⅲ	14	94.37	48.58	96.99	57.23	8.53	4.86	7.12	3.02	105.65	50.65	170.22	81.41

QTN位点 QTN marker	染色体位置 Position	候选基因CandidateGenes	功能注释 Annotation
2_18197441	Chr.2:18197441	GRMZM2G567481	—
2_18739525	Chr.2:18739525	GRMZM2G175504	有丝分裂激活蛋白激酶激酶激酶YODA Mitogen-activated protein kinase kinase kinase YODA
3_155021348	Chr.3:155021348	GRMZM2G146280	含WD重复序列的蛋白质结构域磷酸肌醇相互作用蛋白3 WD repeat-containing protein domain phosphoinositide-interacting protein 3
3_17590250	Chr.3:17590250	GRMZM2G166064	蛋白TRM32 Protein TRM32
3_17805407	Chr.3:17805407	GRMZM2G020941	—
3_17932972	Chr.3:17932972	GRMZM2G100898	DNA甲基化因子1 Factor of DNA methylation 1
4_18511010	Chr.4:18511010	GRMZM5G846314	—
4_28100027	Chr.4:28100027	GRMZM2G319062	多酚氧化酶潜伏型，叶绿体 Polyphenol oxidase latent form, chloroplastic
5_204262820	Chr.5:204262820	GRMZM2G000280	反转录转座子蛋白Ty1-copia亚类 Retrotransposon protein Ty1-copia subclass
9_149806469	Chr.9:149806469	GRMZM2G061314	蛋白结合蛋白 protein binding protein
10_138859255	Chr.10:138859255	GRMZM2G391272	皮层细胞描绘蛋白 Cortical cell-delineating protein