应用东乡野生稻回交重组自交系分析水稻耐低氮产量相关性状QTL

doi:10.16819/j.1001-7216.2020.0408

中国水稻科学 ›› 2020, Vol. 34 ›› Issue (6): 499-511.DOI: 10.16819/j.1001-7216.2020.0408

应用东乡野生稻回交重组自交系分析水稻耐低氮产量相关性状QTL

吴婷¹, 李霞¹, 黄得润², 黄凤林³, 肖宇龙^1,^*(), 胡标林^1,^*()

¹ 江西省农业科学院水稻研究所/水稻国家工程实验室（南昌）,南昌 330200
² 中国水稻研究所国家水稻改良中心/水稻生物学国家重点实验室, 杭州 310006
³ 湖南省水稻研究所农业农村部长江中下游籼稻遗传重点实验室,长沙410125

收稿日期:2020-04-30 修回日期:2020-06-03 出版日期:2020-11-10 发布日期:2020-11-10
通讯作者: 肖宇龙,胡标林
基金资助:
江西省自然科学基金资助项目(20171ACB21071);国家自然科学基金资助项目(31760378);农业农村部长江中下游籼稻遗传育种重点实验室开放课题资助项目(2018KLMA02);水稻生物学国家重点实验室开放课题(20200201)

QTL Analysis for Yield Traits Related to Low Nitrogen Tolerance Using Backcrossing Recombinant Inbred Lines Derived from Dongxiang Wild Rice (Oryza rufipogon Griff.)

Ting WU¹, Xia LI¹, Derun HUANG², Fenglin HUANG³, Yulong XIAO^1,^*, Biaolin HU^1,^*

¹ Rice Research Institute, Jiangxi Academy of Agricultural Sciences / Rice National Engineering Laboratory(Nanchang), Nanchang 330200, China
² Hunan Rice Research Institute/ Key Laboratory of indica Rice Genetics and Breeding in the Middle and Lower Reaches of Yangtze River Valley, Ministry of Agriculture and Rural Affairs, Changsha 410125, China
³ State Key Laboratory of Rice Biology/Chinese National Center for Rice Improvement,China National Rice Research Institute, Hangzhou 310006, China;

Received:2020-04-30 Revised:2020-06-03 Online:2020-11-10 Published:2020-11-10
Contact: Yulong XIAO, Biaolin HU

摘要/Abstract

摘要： 目的东乡野生稻低氮耐性强,是水稻耐低氮育种的重要资源。鉴定东乡野生稻耐低氮基因对研究耐低氮遗传机制、培育耐低氮水稻品种具有重要意义。方法利用协青早B//东乡野生稻/协BBC₁F₁₂回交重组自交系及其遗传图谱,应用Windows QTL Cartographer 2.5分析施氮肥和不施氮肥下的株高和产量相关性状QTL。结果共检测到57个控制株高和产量性状的QTL,分布于10条染色体上的33个区域,单个QTL表型贡献率为3.17%~63.40%,其中32个QTL的增效等位基因来自东乡野生稻。19个QTL在施氮和未施氮条件下均检测到,38个QTL仅在单一环境下检测到显著效应,表明不同施氮水平下水稻性状的遗传机制不同。结论 43个QTL分别聚集于7条染色体上的14个QTL簇,表明不同性状涉及到共同遗传机制,并可通过分子标记辅助选择方法进行耐低氮有利等位基因的聚合育种。

关键词: 耐低氮, 东乡野生稻, 产量性状, 数量性状基因座

Abstract:

【Objective】Dongxiang wildrice (Oryzarufipogon Griff.) has strong low nitrogen toleranceand is an important germplasm for low nitrogen tolerance improvement. Identification of genes responsible for low nitrogen tolerance in Dongxiang wild rice is of great importance to understand molecular mechanisms of low nitrogen tolerance and develop rice varieties with low nitrogen tolerance. 【Method】Quantitative trait loci (QTLs) for plant height and yield traits under low and normal nitrogen conditions wasidentified using backcrossing recombinant inbred lines (BC₁F₁₂) derived from an interspecific cross Xieqingzao B // Dongxiang wild rice/Xieqingzao B and its genetic linkage. 【Result】A total of 57 QTLs were detected in 33 regions on all chromosome, exceptchromosome 4 and 8. They explained individually 3.17%~63.40% phenotypic variation, and 32 QTLsof them had favorable alleles derived from Dongxiang wild rice.NineteenQTLs were simultaneously detected under both nitrogen treatments, and 38QTLs were only identified under single nitrogen treatment, suggesting variousgenetic mechanismsin rice growth and yield formation under low and normal nitrogen conditions. 【Conclusion】Fourteen QTLclusters, 43 QTLs included, scattered on sevenchromosomes, indicatingthe common genetic-physiological mechanisms behind different traits, and theQTL pyramiding for low nitrogen tolerance can be achieved by molecular marker-assisted selection.

Key words: low nitrogen tolerance, Dongxiang wild rice (Oryzarufipogon Griff.), yield trait, quantitative trait locus

中图分类号:

S511.034

吴婷, 李霞, 黄得润, 黄凤林, 肖宇龙, 胡标林. 应用东乡野生稻回交重组自交系分析水稻耐低氮产量相关性状QTL[J]. 中国水稻科学, 2020, 34(6): 499-511.

Ting WU, Xia LI, Derun HUANG, Fenglin HUANG, Yulong XIAO, Biaolin HU. QTL Analysis for Yield Traits Related to Low Nitrogen Tolerance Using Backcrossing Recombinant Inbred Lines Derived from Dongxiang Wild Rice (Oryza rufipogon Griff.)[J]. Chinese Journal OF Rice Science, 2020, 34(6): 499-511.

图/表 5

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性状 Trait	处理 Treatment	亲本Parent		BIL群体 BIL population
性状 Trait	处理 Treatment	东乡野生稻 Dongxiang wild rice	协青早B Xieqingzao B	平均值 Mean	标准差 Standard deviation	变幅 Range	偏度 Skewness	峰度 Kurtosis
穗长 PL / cm	施氮 Nitrogen	25.45^**	18.67	22.46	2.77	15.35~32.16	0.26	0.45
穗长 PL / cm	未施氮Zero-nitrogen	25.51^**	20.66	22.68	2.36	15.72~29.54	0.14	0.04
着粒密度 GD / (No.∙cm^-1)	施氮 Nitrogen	1.92	3.96^**	4.32	1.44	1.18~9.90	1.14	1.91
着粒密度 GD / (No.∙cm^-1)	未施氮Zero-nitrogen	2.26	4.10^**	4.56	1.41	1.95~9.83	1.14	1.91
千粒重 TGW / g	施氮 Nitrogen	20.22	24.90^**	23.36	3.00	15.45~34.57	0.17	0.45
千粒重 TGW / g	未施氮Zero-nitrogen	19.86	25.45^**	23.53	2.88	16.33~31.47	-0.05	0.02
单株产量 PY / g	施氮 Nitrogen	11.30	23.17^**	18.28	6.23	6.06~47.97	1.17	3.29
单株产量 PY / g	未施氮Zero-nitrogen	14.77	16.54	15.92	5.18	6.39~34.18	0.99	1.31
每穗实粒数 FGP	施氮 Nitrogen	40.64	59.89^*	76.20	30.86	15.35~191.50	1.07	1.44
每穗实粒数 FGP	未施氮Zero-nitrogen	44.40	63.14^*	77.89	27.86	28.66~193.14	1.04	1.91
每穗总粒数 TGP	施氮 Nitrogen	48.90	73.90^**	98.34	37.96	25.49~226.13	0.94	0.65
每穗总粒数 TGP	未施氮Zero-nitrogen	56.62	84.81^**	104.18	35.98	42.65~219.06	0.88	0.73
结实率 SF/ %	施氮 Nitrogen	83.30	80.94	77.61	10.23	46.59~94.38	-1.07	0.89
结实率 SF/ %	未施氮Zero-nitrogen	78.87	74.12	75.32	9.99	42.61~91.87	-0.95	0.68
株高 PH / cm	施氮 Nitrogen	178.00^**	86.00	110.25	23.60	73.67~179.33	0.65	-0.40
株高 PH / cm	未施氮Zero-nitrogen	164.83^**	88.17	109.78	21.98	67.00~164.67	0.53	-0.76
有效穗数 EP	施氮 Nitrogen	14.00	15.67	11.07	3.29	4.67~26.00	1.18	2.63
有效穗数 EP	未施氮Zero-nitrogen	16.83^**	10.33	9.19	2.51	4.67~18.67	1.14	1.91

性状 Trait	处理 Treatment	亲本Parent		BIL群体 BIL population
性状 Trait	处理 Treatment	东乡野生稻 Dongxiang wild rice	协青早B Xieqingzao B	平均值 Mean	标准差 Standard deviation	变幅 Range	偏度 Skewness	峰度 Kurtosis
穗长 PL / cm	施氮 Nitrogen	25.45^**	18.67	22.46	2.77	15.35~32.16	0.26	0.45
穗长 PL / cm	未施氮Zero-nitrogen	25.51^**	20.66	22.68	2.36	15.72~29.54	0.14	0.04
着粒密度 GD / (No.∙cm^-1)	施氮 Nitrogen	1.92	3.96^**	4.32	1.44	1.18~9.90	1.14	1.91
着粒密度 GD / (No.∙cm^-1)	未施氮Zero-nitrogen	2.26	4.10^**	4.56	1.41	1.95~9.83	1.14	1.91
千粒重 TGW / g	施氮 Nitrogen	20.22	24.90^**	23.36	3.00	15.45~34.57	0.17	0.45
千粒重 TGW / g	未施氮Zero-nitrogen	19.86	25.45^**	23.53	2.88	16.33~31.47	-0.05	0.02
单株产量 PY / g	施氮 Nitrogen	11.30	23.17^**	18.28	6.23	6.06~47.97	1.17	3.29
单株产量 PY / g	未施氮Zero-nitrogen	14.77	16.54	15.92	5.18	6.39~34.18	0.99	1.31
每穗实粒数 FGP	施氮 Nitrogen	40.64	59.89^*	76.20	30.86	15.35~191.50	1.07	1.44
每穗实粒数 FGP	未施氮Zero-nitrogen	44.40	63.14^*	77.89	27.86	28.66~193.14	1.04	1.91
每穗总粒数 TGP	施氮 Nitrogen	48.90	73.90^**	98.34	37.96	25.49~226.13	0.94	0.65
每穗总粒数 TGP	未施氮Zero-nitrogen	56.62	84.81^**	104.18	35.98	42.65~219.06	0.88	0.73
结实率 SF/ %	施氮 Nitrogen	83.30	80.94	77.61	10.23	46.59~94.38	-1.07	0.89
结实率 SF/ %	未施氮Zero-nitrogen	78.87	74.12	75.32	9.99	42.61~91.87	-0.95	0.68
株高 PH / cm	施氮 Nitrogen	178.00^**	86.00	110.25	23.60	73.67~179.33	0.65	-0.40
株高 PH / cm	未施氮Zero-nitrogen	164.83^**	88.17	109.78	21.98	67.00~164.67	0.53	-0.76
有效穗数 EP	施氮 Nitrogen	14.00	15.67	11.07	3.29	4.67~26.00	1.18	2.63
有效穗数 EP	未施氮Zero-nitrogen	16.83^**	10.33	9.19	2.51	4.67~18.67	1.14	1.91

处理 Treatment	性状 Trait	穗长 PL	着粒密度GD	千粒重TGW	单株产量PY	每穗实粒数FGP	每穗总粒数TGP	结实率SF	株高 PH
未施氮 Zero-nitrogen	着粒密度GD	0.24^**
	千粒重TGW	0.05	-0.41^**
	单株产量PY	0.32^**	0.54^**	0.04
	每穗实粒数FGP	0.52^**	0.85^**	-0.34^**	0.63^**
	每穗总粒数TGP	0.50^**	0.96^**	-0.35^**	0.58^**	0.91^**
	结实率SF	0.09	-0.17^*	0.00	0.21^**	0.28^**	-0.13
	株高PH	0.61^**	-0.03	0.09	0.13	0.27^**	0.15^*	0.31^**
	有效穗数EP	-0.37^**	-0.30^**	0.05	0.31^**	-0.40^**	-0.36^**	-0.11	-0.28^**
施氮 Nitrogen application	着粒密度GD	0.28^**
	千粒重TGW	0.09	-0.41^**
	单株产量PY	0.39^**	0.54^**	-0.03
	每穗实粒数FGP	0.55^**	0.87^**	-0.32^**	0.66^**
	每穗总粒数TGP	0.55^**	0.95^**	-0.32^**	0.60^**	0.93^**
	结实率SF	0.08	-0.08	-0.05	0.27^**	0.30^**	-0.04
	株高PH	0.25^**	0.04	0.01	-0.03	0.14^*	0.11	0.08
	有效穗数EP	-0.37^**	-0.36^**	-0.05	0.22^**	-0.43^**	-0.42^**	-0.12	-0.19^**

处理 Treatment	性状 Trait	穗长 PL	着粒密度GD	千粒重TGW	单株产量PY	每穗实粒数FGP	每穗总粒数TGP	结实率SF	株高 PH
未施氮 Zero-nitrogen	着粒密度GD	0.24^**
	千粒重TGW	0.05	-0.41^**
	单株产量PY	0.32^**	0.54^**	0.04
	每穗实粒数FGP	0.52^**	0.85^**	-0.34^**	0.63^**
	每穗总粒数TGP	0.50^**	0.96^**	-0.35^**	0.58^**	0.91^**
	结实率SF	0.09	-0.17^*	0.00	0.21^**	0.28^**	-0.13
	株高PH	0.61^**	-0.03	0.09	0.13	0.27^**	0.15^*	0.31^**
	有效穗数EP	-0.37^**	-0.30^**	0.05	0.31^**	-0.40^**	-0.36^**	-0.11	-0.28^**
施氮 Nitrogen application	着粒密度GD	0.28^**
	千粒重TGW	0.09	-0.41^**
	单株产量PY	0.39^**	0.54^**	-0.03
	每穗实粒数FGP	0.55^**	0.87^**	-0.32^**	0.66^**
	每穗总粒数TGP	0.55^**	0.95^**	-0.32^**	0.60^**	0.93^**
	结实率SF	0.08	-0.08	-0.05	0.27^**	0.30^**	-0.04
	株高PH	0.25^**	0.04	0.01	-0.03	0.14^*	0.11	0.08
	有效穗数EP	-0.37^**	-0.36^**	-0.05	0.22^**	-0.43^**	-0.42^**	-0.12	-0.19^**

性状 Trait	处理 Treatment	数量性状位点 QTL	标记区间 Interval		加性效应^a Additive effect^a	贡献率 Proportion of the variance explained/%	文献 Reference
穗长PL	未施氮 Zero-nitrogen	qPL1	RM212-InDel 1	5.42	1.14	11.54
		qPL5	RM164-RM188	6.65	1.49	21.13
		qPL6	RM3805-RM204	4.12	-0.86	6.66	[13]
		qPL10	RM1375-RM5620	4.19	-1.00	10.24
	施氮 Nitrogen	qPL1	InDel 1-RM6840	6.89	1.20	16.39
		qPL5	RM164-RM188	9.93	1.58	32.46
		qPL10	RM1375-RM5620	4.33	-0.89	10.72
着粒密度 GD	未施氮 Zero-nitrogen	qGD1.1	RM10176-RM243	13.92	0.87	16.88	[36]
		qGD1.2	RM3475-RM246	3.05	-0.31	3.23
		qGD3	RM6266-RM168	2.75	0.94	14.38	[13]
		qGD5	RM289-RM249	2.79	0.84	18.79
		qGD6	RM508-RM3805	3.25	-0.37	5.19
		qGD9	RM553-RM160	7.13	1.07	19.52
		qGD11	RM224-RM144	7.51	0.63	11.47
		qGD12	RM235-RM17	3.34	0.49	5.67
	施氮 Nitrogen	qGD1.1	RM10176-RM243	10.97	0.76	12.60	[36]
		qGD1.2	RM3475-RM246	4.16	-0.35	4.24
		qGD2	RM29-RM475	3.14	-0.38	4.30
		qGD6	RM508-RM3805	2.50	-0.39	5.06
		qGD9	RM1896-RM553	3.06	0.46	5.41
		qGD11	RM224-RM144	5.17	0.48	6.90
千粒重 TGW	未施氮 Zero-nitrogen	qTGW1	RM129-RM3475	2.15	0.82	5.05	[44]
		qTGW6	RM253-RM276	2.87	1.05	9.65
		qTGW7	RM481-InDel7.1	3.63	-1.40	8.12
		qTGW10	RM304-RM171	5.46	-1.05	10.36
	施氮 Nitrogen	qTGW2	RM262-RM13637	2.44	1.13	6.76
		qTGW3	RM282-RM16	2.48	-0.78	5.61
		qTGW7	RM481-InDel7.1	3.12	-1.49	8.53
		qTGW10	RM171-RM590	4.42	-0.95	9.67
		qTGW11	RM286-RM332	4.15	-1.77	6.49
单株产量 PY	未施氮 Zero-nitrogen	qPY3	RM22-RM569	3.27	-2.29	5.88
	未施氮 Zero-nitrogen	qPY5.1	RM188-RM3870	4.04	2.09	9.21	[39]
	施氮 Nitrogen	qPY5.2	RM289-RM249	3.07	3.10	19.14
		qPY5.1	RM188-RM3870	4.55	2.29	14.92	[39]
		qPY9.1	InDel 9.1-RM444	3.57	-1.87	6.28
		qPY9.2	RM1896-RM553	4.38	2.85	13.94
每穗实粒数 FGP	未施氮 Non-nitrogen	qFGP1	RM10176-RM243	9.48	22.22	27.09	[13, 35]
	Zero-nitrogen	qFGP2	RM71-RM300	3.71	-9.39	6.65	[13, 35, 38]
		qFGP5	RM188-RM3870	5.55	13.39	13.50
		qFGP6	RM3805-RM204	3.30	-7.95	5.04
		qFGP9	RM553-RM160	4.29	23.93	21.89
		qFGP11	RM224-RM144	3.28	12.49	10.10
	施氮 Nitrogen	qFGP1	RM10176-RM243	7.91	17.96	23.08	[13, 35]
	Nitrogen	qFGP5	RM188-RM3870	3.10	9.39	8.04
		qFGP9	RM553-RM160	3.07	-8.87	5.28
		qFGP10	RM1375-RM5620	2.69	9.02	5.43

应用东乡野生稻回交重组自交系分析水稻耐低氮产量相关性状QTL

QTL Analysis for Yield Traits Related to Low Nitrogen Tolerance Using Backcrossing Recombinant Inbred Lines Derived from Dongxiang Wild Rice (Oryza rufipogon Griff.)

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Metrics

性状 Trait	处理 Treatment	数量性状位点 QTL	区间 Interval	LOD值 LOD value	加性效应^a Additiveeffect^a	贡献率 Proportion of the variance explained/%
穗总粒数 TGP	未施氮 Zero-nitrogen	qTGP1	RM10176-RM243	12.90	27.76	28.44	[13, 35, 36]
		qTGP2	RM5870-RM71	3.27	-10.23	5.00	[13, 35, 38]
		qTGP5	RM188-RM3870	7.26	17.90	15.75
		qTGP6	RM3805-RM204	4.64	-10.74	6.11
		qTGP9.2	RM553-RM160	4.85	26.36	16.73
		qTGP10	RM1375-RM5620	2.25	-8.56	4.46
		qTGP11	RM224-RM144	4.39	16.59	11.96
	施氮 Nitrogen	qTGP1	RM10176-RM243	11.96	23.46	20.22	[13, 35, 36]
		qTGP5	RM188-RM3870	2.92	10.33	5.96
		qTGP9.1	InDel 9.1-RM444	3.33	16.31	8.57
		qTGP9.2	RM1896-RM553	3.43	-10.37	7.34
结实率 SF	未施氮 Zero-nitrogen	qSF1.1	RM572-RM140	3.24	3.80	7.67
		qSF2	RM213-RM208	3.34	-8.92	36.69
		qSF6	RM253-RM276	2.54	3.09	7.26
		qSF7	RM432-RM11	2.80	-5.03	8.06
		qSF12	RM20B-RM19	3.85	-10.16	37.01
	施氮 Nitrogen	qSF1.1	RM572-RM140	3.06	3.20	6.10
		qSF1.2	RM212-InDel 1	2.77	2.49	5.41
		qSF3	RM7-RM218	4.55	-8.88	27.21
		qSF9	InDel 9.1-RM444	3.70	-9.65	21.43
		qSF12	RM20B-RM19	4.09	-9.54	37.99
株高 PH	未施氮 Zero-nitrogen	qPH1.2	RM572-RM140	2.15	5.32	3.17
		qPH1.3	RM212-InDel1	22.01	18.92	57.22	[36, 37, 40]
		qPH6	RM204-RM225	3.20	-5.49	4.41
	施氮 Nitrogen	qPH1.1	RM10176-RM243	4.34	23.13	63.40
有效穗数 EP	未施氮 Zero-nitrogen	qEP9	InDel9.1-RM444	3.29	3.00	33.59
	施氮 Nitrogen	qEP1	InDel1-RM6840	2.03	-0.90	5.56
		qEP3.1	RM22-RM569	2.07	0.95	4.07
		qEP3.2	RM168-RM85	3.49	2.85	28.69
		qEP6	InDel6.1-InDel6.2	2.80	3.57	27.10
^a加性效应是指东乡野生稻等位基因取代协青早B等位基因。 ^aAdditive effect of replacing an Xieqingzao B allele by Dongxiang wild rice allele.

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