基于图像分析的玉米抗拟轮枝镰孢穗腐病的QTL定位

doi:10.3864/j.issn.0578-1752.2021.13.003

摘要/Abstract

摘要：

【目的】玉米穗腐病是一种在全世界广泛发生且危害严重的真菌性病害,其中,拟轮枝镰孢引起的穗腐病（Fusarium ear rot,FER）在中国发生最为普遍。通过图像分析方法进行FER抗病QTL定位,并对前期通过病害评级方法定位的FER抗病QTL进行验证,探索一种新的玉米穗腐病的病害鉴定方法,为玉米穗腐病的遗传改良提供依据。【方法】利用高感FER的自交系（ZW18）分别与3个高抗自交系（承351、丹598和吉V203）构建F₂群体（F₂-C、F₂-D和F₂-J）和相应的F_2﹕3家系,通过图像分析的方法获得每个F_2﹕3家系的果穗病斑百分比,进而定位玉米FER抗病QTL。【结果】3个群体共定位到18个FER抗病QTL,其中,分别位于2.02—2.03 bins、4.06—4.07 bins和8.06 bin上的3个QTL（qRf2、qRf3和qRf4）可解释的表型变异率分别高达21.80%、25.80%和27.40%。F₂-J群体的qRf11与F₂-C群体的qRf1和F₂-D群体的qRf6在第1染色体均有重叠区间,可解释的表型变异率达到16.50%。来自F₂-D群体的qRf9与F₂-J群体的qRf16在第8染色体8.05 bin有重叠区间,且抗性基因均来源于抗性亲本。F₂-C群体的qRf3与F₂-J群体的qRf15在第4染色体4.06—4.07 bins有重叠区间。另外,与之前通过病害评级方法定位的结果相比,qRf1、qRf6和qRf11在1.06—1.07 bins与评级方法定位的抗病位点qRfer13重合,qRf3和qRf15在4.06—4.07 bins与评级方法定位的抗病位点qRfer3和qRfer17重叠,qRf7与qRfer6在2.04 bin的定位区间完全重合,qRf17与qRfer20在S2重复中定位到9.03—9.05 bins的重叠区间,且来源于相同的抗源。【结论】定位到18个FER抗性位点,其中,位于1.04—1.07 bins、4.06—4.07 bins和8.05 bin上的抗病位点在不同群体中均可以被检测到,位于2.04 bin和9.03—9.05 bins上的抗病位点用不同的检测方法可以被检测到,表明在这些区间可能存在FER的抗性位点。QTL的定位区间在不同群体中的重叠性在一定程度上验证了定位区间的真实性,不同方法之间定位到重叠区间,说明利用图像分析方法定位FER抗病QTL具有一定的准确性。

关键词: 玉米, 拟轮枝镰孢穗腐病, 抗性, QTL

Abstract:

【Objective】Ear rot in maize is a kind of fungal disease that is prevalent and giving serious threat to maize production worldwide, among which,Fusarium ear rot (FER) is the most commonly reported in China. The location of FER disease resistance QTL is very important for the genetic improvement of maize ear rot. Locating FER disease-resistant QTL by an image analysis method not only explores a new disease evaluation method for FER in maize, but also validated the FER resistance QTLs located in the early stage through the disease rating method. 【Method】In this study, three F₂ populations (namely F₂-C, F₂-D and F₂-J) and their corresponding F_2:3 families were used for QTL mapping of resistance to FER, which were constructed from a highly FER-susceptible inbred line ZW18 and three highly FER-resistant inbred lines (Cheng 351, Dan 598 and JiV203) respectively. The percentage of diseased spots was obtained by image analysis for the ears of F_2:3 families, then the FER resistance QTLs were mapped.【Result】18 FER resistance QTLs were mapped. Among them, qRf2,qRf3 and qRf4 located in 2.02-2.03 bins, 4.06-4.07 bins and 8.06 bin explained the phenotypic variation as high as 21.80%, 25.80% and 27.40%, respectively. The qRf11 of the F₂-J population and the qRf1 of the F₂-C population and the qRf6 of the F₂-D population all have an overlapping interval, and the explainable phenotypic variation rate reached 16.50%. The qRf9 from the F₂-D population and theqRf16 from the F₂-J population have an overlapping interval in 8.05 bin, and the qRf3 from the F2-C population and the qRf15 from the F₂-J population have an overlapping interval in the 4.06-4.07 bins of 4^th chromosome. In addition, compared with the results of the previous positioning by the disease rating method,qRf1, qRf6 andqRf11 overlap with the disease resistance site qRfer13 located by the rating method at 1.06-1.07 bins, and qRf3 and qRf15 at 4.06-4.07 bins and the resistance location located by the rating method. The disease site qRfer3 and qRfer17overlap, qRf7 and qRfer6 completely overlap in the 2.04 bin location interval, qRf17 and qRfer20 are located in the overlap interval of 9.03-9.05 bins in the S2 duplication, and are derived from the same source of resistance. 【Conclusion】 The 18 FER resistance sites located in 3 populations, among which the resistance sites located in 1.04-1.07 bins, 4.06-4.07 bins and 8.05 bin can be detected in different populations, located in 2.04 bin and 9.03-9.05 bins can be detected by different detection methods, indicating that FER resistance sites may exist in these intervals. The overlap of QTL positioning intervals in different populations verifies the authenticity of the positioning intervals to a certain extent. The overlapping intervals are located between different methods, indicating that the image analysis method is used to locate FER disease-resistant QTLs with certain accuracy.

Key words: maize, Fusarium verticilloides, resistance, QTL

闻竞,沈彦岐,王梓钰,李世界,莫蓝月,雷宇豪,张艳,韩四平. 基于图像分析的玉米抗拟轮枝镰孢穗腐病的QTL定位[J]. 中国农业科学, 2021, 54(13): 2724-2736.

WEN Jing,SHEN YanQi,WANG ZiYu,LI ShiJie,MO LanYue,LEI YuHao,ZHANG Yan,HAN SiPing. QTL Mapping of Resistance to Fusarium Ear Rot in Maize Based on Image Analysis[J]. Scientia Agricultura Sinica, 2021, 54(13): 2724-2736.

图/表 9

图1

图2

图3

表1

表2

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图5

表3

玉米FER抗病QTLs及其参数"

位点 QTL	染色体 Chr.	位置Site (Bins)	重复 Replication	侧翼标记 Flanking markers	LOD	加性效应 Additive	显性效应 Dominance	抗性来源 SRA	表型变异率 R²(%)
qRf1	1	1.04—1.06	BLUE	e1583803—e1188929	4.78	1.6956	-0.7136	承351 Cheng351	15.60
qRf2	2	2.02—2.03	S1	e2102115—e2161221	3.84	1.6043	-1.4111	承351 Cheng351	21.80
	2	2.02	S	e2102115—e2126291	3.78	1.5500	-2.0277	承351 Cheng351	20.30
	2	2.02—2.03	BLUE	e2102115—e2161221	4.49	1.9089	0.3621	承351 Cheng351	9.70
qRf3	4	4.06—4.07	G1	e4156956—e4175573	6.05	3.7277	-2.9966	承351 Cheng351	25.80
qRf3	4	4.06—4.07	BLUE	e4167183—e4179848	5.41	2.0177	-0.8886	承351 Cheng351	19.60
qRf4	8	8.06	G1	e8161042—e8165169	5.61	3.9752	-3.3116	承351 Cheng351	27.40
qRf4	8	8.06	BLUE	e8153595—e8165169	6.38	2.3975	-1.2185	承351 Cheng351	26.00
qRf5	8	8.07—8.08	BLUE	e8167666—e8171604	4.19	-2.0687	-0.6555	ZW18	7.10
qRf6	1	1.07	BLUE	e1197737—e1212331	4.16	-1.3688	1.6416	ZW18	10.82
qRf7	2	2.04	BLUE	e2365995—e2639635	5.93	2.1626	0.9592	丹598 Dan598	7.94
qRf8	3	3.03—3.04	S	e3106087—e3335621	4.09	2.1894	-1.1950	丹598 Dan598	10.04
qRf9	8	8.05—8.06	BLUE	e8130926—e8150778	4.48	1.8506	-0.6254	丹598 Dan598	10.85
qRf10	10	10.06	S	e1013859—e1014290	4.17	2.4951	-1.4593	丹598 Dan598	12.32
qRf11	1	1.06—1.07	S2	e1178476—e1200620	5.69	2.8361	-1.3421	吉V203 JiV203	16.50
qRf12	3	3.04—3.05	S1	e3170341—e3154562	4.08	-2.9888	0.8469	ZW18	13.46
qRf13	3	3.06—3.07	G1	e3180570—e3195935	3.91	-4.3654	-0.5613	ZW18	7.38
qRf14	4	4.01—4.02	G1	e4359518—e4107296	3.69	3.4943	-2.2933	吉V203 JiV203	11.93
qRf14	4	4.01—4.03	BLUE	e4359518—e4120869	4.56	1.8443	-0.5136	吉V203 JiV203	11.65
qRf15	4	4.05—4.07	G2	e4689700—e4171651	5.91	30.334	-0.8452	吉V203 JiV203	17.96
qRf15	4	4.06—4.07	S2	e4168742—e4178823	5.25	3.4990	0.6582	吉V203 JiV203	8.16
qRf16	8	8.03—8.05	S	e8842299—e8144487	4.59	3.5113	-2.2912	吉V203 JiV203	8.15
qRf17	9	9.03—9.05	S2	e9432770—e9128660	5.55	-2.5901	-2.3177	ZW18	3.61
qRf18	9	9.06—9.07	G	e9143806—e9149382	3.93	-2.2164	-1.6562	ZW18	2.85

表3

表4

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群体 Populations	重复 Replication	抗病亲本 Resistant parents	感病亲本 Susceptible parents	F₂群体
群体 Populations	重复 Replication	抗病亲本 Resistant parents	感病亲本 Susceptible parents	平均值 Mean	标准差 SD	偏度 Skewness	峰度 Kurtosis	变异系数 CV (%)	遗传力 H²
F₂-C	G1	0.0	49.22	6.24	7.09	3.38	15.16	113.63	0.63
	G2	0.0	82.26	5.14	4.70	2.50	11.67	91.46
	S1	0.0	63.85	1.65	3.06	5.30	33.16	185.50
	S2	0.0	55.38	5.41	5.09	1.96	6.41	94.03
F₂-D	G1	0.0	49.22	8.30	8.80	5.60	51.70	106.03	0.29
	G2	0.0	82.26	7.75	5.54	1.16	2.04	71.50
	S1	0.0	63.85	6.11	8.62	6.61	67.02	141.03
	S2	2.4	55.38	7.03	8.52	6.53	66.75	121.22
F₂-J	G1	0.0	49.22	9.74	9.30	5.60	48.80	95.52	0.56
	G2	0.0	82.26	7.73	5.34	1.23	1.51	69.02
	S1	1.3	63.85	4.72	4.78	2.44	8.14	101.11
	S2	0.0	55.38	7.86	6.03	1.39	2.45	76.72

群体 Populations	变异来源 Sources	自由度 Df	离均差平方和 SS	均方 MS	F值 F value	P值 P value (>F_0.001)	显著性 Significance	偏度分析 Pr
F₂-C	G	117	5803.8	49.60	2.1406	8.118e-07	***
	L	1	553.7	553.67	23.8927	2.015e-06	***
	R	1	179.1	179.09	7.7283	0.005928	**
	G:L	107	1952.0	18.24	0.7873	0.916935
	Residuals	211	4889.6	23.17				7.410274e-15
F₂-D	G	199	17047.6	85.67	1.5483	0.0001392	***
	L	1	410.4	410.45	7.4184	0.0067446	**
	R	1	7.5	7.55	0.1364	0.7120661
	G:L	196	11814.3	60.28	1.0894	0.2392358
	Residuals	392	21688.7	55.33				1.390253e-14
F₂-J	G	173	12665.6	73.21	1.8950	3.547e-07	***
	L	1	941.4	941.42	24.3679	1.261e-06	***
	R	1	41.6	41.61	1.0770	0.3001
	G:L	165	5293.4	32.08	0.8304	0.9113
	Residuals	333	12864.9	38.63				-8.235978e-13

位点 Name	染色体 Chr.	位置 Site (Bins)	重复 Replication	侧翼标记 Flanking markers	LOD	加性效应 Additive	显性效应 Dominant	抗性来源 SRA	表型变异率 R²(%)
qRfer3	4	4.06—4.07	G	e4163306—e4177040	4.22	0.5365	-0.2592	承351 Cheng351	18.34
qRfer3	4	4.06—4.08	BLUE	e4167183—e4179848	4.78	0.4623	-0.1894	承351 Cheng351	17.76
qRfer6	2	2.04	BLUE	e2365995—e2639635	5.12	0.3268	0.0293	丹598 Dan598	7.02
qRfer13	1	1.05—1.06	S1	e1100903—e1182999	5.13	0.3924	0.2506	吉V203 JiV203	5.45
qRfer13	1	1.06—1.07	S2	e1178476—e1202995	7.40	0.6921	-0.1173	吉V203 JiV203	16.73
qRfer17	4	4.05—4.06	G1	e4327859—e4163646	5.97	0.6190	-0.0758	吉V203 JiV203	12.02
		4.05—4.07	G2	e4978482—e4172059	7.83	0.7965	-0.3242	吉V203 JiV203	21.81
		4.05—4.06	G	e4327859—e4163646	8.42	0.6540	-0.1231	吉V203 JiV203	17.96
		4.05—4.06	BLUE	e4327859—e4163646	6.65	0.5549	-0.0784	吉V203 JiV203	17.37
qRfer20	9	9.03—9.05	S2	e9432770—e9130925	6.81	-0.3540	-0.4352	ZW18	4.10