辣椒雄性不育分子研究进展

doi:10.11924/j.issn.1000-6850.casb2023-0860

中国农学通报 ›› 2024, Vol. 40 ›› Issue (13): 27-35.doi: 10.11924/j.issn.1000-6850.casb2023-0860

辣椒雄性不育分子研究进展

徐晓美¹(), 衡周¹, 王恒明¹, 孙启迪², 徐小万¹()

¹ 广东省农业科学院蔬菜研究所/广东省蔬菜新技术研究重点实验室，广州 510640
² 茂名市茂蔬种业科技有限公司，广东茂名 525000

收稿日期:2023-12-14 修回日期:2024-02-27 出版日期:2024-04-28 发布日期:2024-04-28
通讯作者:
徐小万，男，1975年出生，湖南郴州人，研究员，博士，研究方向为茄果类蔬菜遗传育种。通信地址：530460 广东省广州市天河区五山镇金颖路66号蔬菜研究所，Tel：020-38469456，E-mail：xxw7505@163.com。
作者简介:
徐晓美，女，1983年出生，江西吉安人，研究员，博士，研究方向为辣椒雄性不育及分子育种研究。通信地址：530460 广东省广州市天河区五山镇金颖路66号蔬菜研究所，Tel：020-38469456，E-mail：xiaomeixu@gdaas.cn。
基金资助:
广东省基础与应用基础研究基金项目“辣椒根腐疫病候选抗性基因功能验证及其抗病分子机制研究”(2021A1515012132); 广东省基础与应用基础研究基金（粤桂联合基金）项目“辣椒根腐疫病候选抗性基因CaRLK功能验证及其与Avr蛋白互作的分子机制研究”(2020A1515410005); 2022年省级乡村振兴战略专项资金种业振兴项目“瓜果类蔬菜核心种质指纹图谱构建”(2022-NJS-00-005); 广州市科技计划项目“多技术融合选育华南特色辣椒新品种”(2023B03J1082); 广东省农业厅种业振兴与科技兴农项目“高产优质辣椒新品种选育研究”(2022-NPY-00-024)

Molecular Advancements of Male Sterility in Pepper

XU Xiaomei¹(), HENG Zhou¹, WANG Hengming¹, SUN Qidi², XU Xiaowan¹()

¹ Vegetable Research Institute, Guangdong Academy of Agriculture Sciences/Guangdong Key Laboratory for New Technology Research of Vegetables, Guangzhou 510640
² Maoming Maoshu Seed Technology Co., Ltd., Maoming, Guangdong 525000

Received:2023-12-14 Revised:2024-02-27 Published:2024-04-28 Online:2024-04-28

摘要/Abstract

摘要：

为对辣椒雄性不育分子研究取得的进展有更直观、系统、深入的了解，梳理已鉴定的辣椒核雄性不育(NMS)基因及部分NMS基因分子标记开发，重点阐述近年来核雄性不育基因精细定位及候选基因探究工作；同时回顾辣椒胞质雄性不育(CMS)相关的线粒体不育基因及其关联分子标记和CMS恢复基因分子标记开发，重点概述近年来恢复基因精细定位研究取得的进展。研究指出，精细定位NMS基因，明确候选基因并开发功能标记是辣椒NMS研究的重点，既能为探明辣椒NMS机理奠定基础又可为育种提供高效利用的分子标记。而CMS相较于NMS更为复杂，表现在遗传机制多样且更易受环境影响等方面，因此，从对CMS育性恢复基因的精细定位到基因克隆以及功能研究并探明CMS分子机制还有很长的一段路要走。

关键词: 辣椒, 核雄性不育, 胞质雄性不育, 恢复基因, 分子标记

Abstract:

In order to have a more intuitive, systematic and in-depth understanding of the progress made in molecular research on male sterility in pepper, we sorted out the identified nuclear male sterility (NMS) genes in pepper and the development of molecular markers for some of the NMS genes, and highlighted the fine mapping of the NMS genes and the exploration of the candidate genes in recent years. Meanwhile, we reviewed the mitochondrial sterile genes related to cytoplasmic male sterility (CMS) and their associated molecular markers, as well as molecular markers developed for CMS fertility restoration genes, with a focus on the progress made in recent years in fine mapping of fertility restoration genes. Finally, it was pointed out that the fine mapping of NMS genes, identification of candidate genes and development of functional markers were the main focuses of NMS research in pepper, which could not only lay the foundation for the exploration of the mechanism of NMS in pepper, but also provided molecular markers for efficient use in breeding. Compared with NMS, CMS is more complex, characterized by diverse genetic mechanisms and more susceptible to environmental influences. Therefore, there is still a long way to go from the fine mapping of CMS restorer-of-fertility genes to the cloning of genes and the functional studies to understand the molecular mechanisms of CMS.

Key words: pepper, nuclear male sterility, cytoplasmic male sterility, fertility restoration gene, molecular marker

徐晓美, 衡周, 王恒明, 孙启迪, 徐小万. 辣椒雄性不育分子研究进展[J]. 中国农学通报, 2024, 40(13): 27-35.

XU Xiaomei, HENG Zhou, WANG Hengming, SUN Qidi, XU Xiaowan. Molecular Advancements of Male Sterility in Pepper[J]. Chinese Agricultural Science Bulletin, 2024, 40(13): 27-35.

图/表 5

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基因名称	遗传模式	基因间关系	基因来源	参考文献
ms1	隐性单基因遗传	与ms_w、msc-2互为等位基因，与ms6-ms8、msk非等位，其他未知	‘All Big’自发突变	[6]
ms2	隐性单基因遗传	与ms10、msk互为等位基因，与ms6-ms8非等位，其他未知	‘California Wonder’自发突变	[7]
ms3	隐性单基因遗传	与ms6-ms8、msk、ms_w非等位，其他未知	‘Pasardjishka Kapia794’辐射诱变	[8-9]
ms4	隐性单基因遗传	与ms6-ms8非等位，其他未知	‘Pasardjishka Kapia794’辐射诱变	[8-9]
ms6	隐性单基因遗传	与ms1-ms4非等位，其他未知	‘Zlaten Medal’辐射诱变	[8-9]
ms7	隐性单基因遗传	与ms1-ms4非等位，其他未知	‘Zlaten Medal’辐射诱变	[8-9]
ms8	隐性单基因遗传	与ms1-ms4非等位，其他未知	‘Zlaten Medal’辐射诱变	[8-9]
ms9	隐性单基因遗传	未知	γ射线诱变	[8-9]
ms10(mc-509)	隐性单基因遗传	与ms2、msk互为等位基因	EMS诱变	[8-9]
ms11(mc-705)	隐性单基因遗传	未知	EMS诱变	[8-9]
ms12	隐性单基因遗传	未知	‘Gambo’自发突变	[11]
ms13(ms)	隐性单基因遗传	未知	‘CA452-1’自发突变	[12]
ms14	隐性单基因遗传	未知	‘Kalyanpur selection’自发突变	[13]
ms15	隐性单基因遗传	未知	‘CA-960’自发突变	[12]
Dms	显性单基因遗传	ms5基因突变	自发突变	[8-9]
msk	隐性单基因遗传	与ms2、ms10互为等位基因	自发突变	[14]
ms_w	隐性单基因遗传	与ms1、msc-2互为等位基因	自发突变	[15]
fms	隐性单基因遗传	未知	自发突变	[19]
msc1	隐性单基因遗传	未知	自发突变	[16-17]
msc2	隐性单基因遗传	未知	自发突变	[17-18]
msc-1	隐性单基因遗传	未知	‘沈椒4号’自发突变	[20]
msc-2	隐性单基因遗传	与ms1、ms_w互为等位基因	‘冀研16号’自发突变	[21]
msc-3	隐性单基因遗传	未知	自发突变	[22]

基因名称	遗传模式	基因间关系	基因来源	参考文献
ms1	隐性单基因遗传	与ms_w、msc-2互为等位基因，与ms6-ms8、msk非等位，其他未知	‘All Big’自发突变	[6]
ms2	隐性单基因遗传	与ms10、msk互为等位基因，与ms6-ms8非等位，其他未知	‘California Wonder’自发突变	[7]
ms3	隐性单基因遗传	与ms6-ms8、msk、ms_w非等位，其他未知	‘Pasardjishka Kapia794’辐射诱变	[8-9]
ms4	隐性单基因遗传	与ms6-ms8非等位，其他未知	‘Pasardjishka Kapia794’辐射诱变	[8-9]
ms6	隐性单基因遗传	与ms1-ms4非等位，其他未知	‘Zlaten Medal’辐射诱变	[8-9]
ms7	隐性单基因遗传	与ms1-ms4非等位，其他未知	‘Zlaten Medal’辐射诱变	[8-9]
ms8	隐性单基因遗传	与ms1-ms4非等位，其他未知	‘Zlaten Medal’辐射诱变	[8-9]
ms9	隐性单基因遗传	未知	γ射线诱变	[8-9]
ms10(mc-509)	隐性单基因遗传	与ms2、msk互为等位基因	EMS诱变	[8-9]
ms11(mc-705)	隐性单基因遗传	未知	EMS诱变	[8-9]
ms12	隐性单基因遗传	未知	‘Gambo’自发突变	[11]
ms13(ms)	隐性单基因遗传	未知	‘CA452-1’自发突变	[12]
ms14	隐性单基因遗传	未知	‘Kalyanpur selection’自发突变	[13]
ms15	隐性单基因遗传	未知	‘CA-960’自发突变	[12]
Dms	显性单基因遗传	ms5基因突变	自发突变	[8-9]
msk	隐性单基因遗传	与ms2、ms10互为等位基因	自发突变	[14]
ms_w	隐性单基因遗传	与ms1、msc-2互为等位基因	自发突变	[15]
fms	隐性单基因遗传	未知	自发突变	[19]
msc1	隐性单基因遗传	未知	自发突变	[16-17]
msc2	隐性单基因遗传	未知	自发突变	[17-18]
msc-1	隐性单基因遗传	未知	‘沈椒4号’自发突变	[20]
msc-2	隐性单基因遗传	与ms1、ms_w互为等位基因	‘冀研16号’自发突变	[21]
msc-3	隐性单基因遗传	未知	自发突变	[22]

标记类型	标记名称	引物序列		NMS 基因	距离/ cM	染色体定位	参考文献
标记类型	标记名称	前引物(5’-3’)	后引物(5’-3’)	NMS 基因	距离/ cM	染色体定位	参考文献
CAPS	PmsM1-CAPS	TGGGCTATCCCGTAAGCCAGCTCAT	GCCGAATCCCTTCATCCTATTTCTCCT	ms1	2~3	-	[25]
SCAR	MS1-SCAR	-	-	ms1	3	-	[26]
HRM	32187928-HRM	CAAAAGTCCATGATAGAGTGACCA	GTATCAATGGCTCATTGGAGATT	ms1	0	5	[27]
CAPS	GMS3-CAPS	-	-	ms3	-	-	[28]
CAPS	HPGMS2CAPs	GGTACTTTGACCCTCATATTGG	TTGTTTGTGGTGTACGTGCT	ms3	3.83	1	[15]
dCAPS	HPGMS3DCAPs	GGGATGTTCAGTCTCATGT	GACTTTTTCCCGATCTCGG	ms3	3.83	1	[15]
SCAR	SCAR_{_}N2	ATACCCAAATCCCACCGTTCA	AATAGGATCAAACTTCGACCAGAAA	ms8	16.1	4	[29]
SCAR	SCAR_V01	ACATGGCCTGAGATCGTGAA	TGTAACCATCTCCCAAATAGAGC	ms8	18.1	4	[29]
SCAR	SCAR_P2	AAGCCAACAGGGGGTATCGCATAAGCA	GGAAGCCAACAACACCCATATTTTCCA	ms8	4.6	4	[29]
SCAR	SCAR_V17	ACCGGCTTGTCCCCGTGGA	ACCGGCTTGTATGACTCTCTA	ms8	6.8	4	[29]
SSR	AVRDC_PP12	TCCTAACTCTTCCCACCACC	GGAGAAGTGTAGCTCCAGCC	ms10	7.2	1	[30]
SSR	AVRDC_MD997	CTGTTTGGAAGGAACCTTTA	TCAGTGATCCAATAATGTGC	ms10	20.8	1	[30]
CAPS	GMSK-CAPS	-	-	msk	0	-	[28]
dCAPS	SPGMS1	TGCAGAAATGGATC	AGGATTTACAAGGAGATGAA	ms_w	0	5	[15]
InDel	Indel-12	GCCTGAAGTTCAAGTGGCTC	CCACAGATGCAGTAACAAGCA	msc-1	0	2	[20]
dCAPS	dCAPs-msc-2	TTGTATCAATGGCTCATTGGAGATTGCGTCT	AACTCCCTCCCTGGCTACAAATATGTCC	msc-2	0	5	[21]
InDel	Ind198	TTCTCCTCCTGCTGTGCATG	TCTCCACCACAAGTCCCTCA	msc-3	0	10	[22]
SSR	H12	GTTCTATTAAGGGTGGTGAGGGT	CTCGCAGCAGATGGTAGTTTAGT	msc2	0.29	-	[32]
SSR	H24	TTAAACACGGGATGCAGCTACTA	AAGTTCCTTTTCGTGTAACGACC	msc2	0	-	[32]

标记类型	标记名称	引物序列		NMS 基因	距离/ cM	染色体定位	参考文献
标记类型	标记名称	前引物(5’-3’)	后引物(5’-3’)	NMS 基因	距离/ cM	染色体定位	参考文献
CAPS	PmsM1-CAPS	TGGGCTATCCCGTAAGCCAGCTCAT	GCCGAATCCCTTCATCCTATTTCTCCT	ms1	2~3	-	[25]
SCAR	MS1-SCAR	-	-	ms1	3	-	[26]
HRM	32187928-HRM	CAAAAGTCCATGATAGAGTGACCA	GTATCAATGGCTCATTGGAGATT	ms1	0	5	[27]
CAPS	GMS3-CAPS	-	-	ms3	-	-	[28]
CAPS	HPGMS2CAPs	GGTACTTTGACCCTCATATTGG	TTGTTTGTGGTGTACGTGCT	ms3	3.83	1	[15]
dCAPS	HPGMS3DCAPs	GGGATGTTCAGTCTCATGT	GACTTTTTCCCGATCTCGG	ms3	3.83	1	[15]
SCAR	SCAR_{_}N2	ATACCCAAATCCCACCGTTCA	AATAGGATCAAACTTCGACCAGAAA	ms8	16.1	4	[29]
SCAR	SCAR_V01	ACATGGCCTGAGATCGTGAA	TGTAACCATCTCCCAAATAGAGC	ms8	18.1	4	[29]
SCAR	SCAR_P2	AAGCCAACAGGGGGTATCGCATAAGCA	GGAAGCCAACAACACCCATATTTTCCA	ms8	4.6	4	[29]
SCAR	SCAR_V17	ACCGGCTTGTCCCCGTGGA	ACCGGCTTGTATGACTCTCTA	ms8	6.8	4	[29]
SSR	AVRDC_PP12	TCCTAACTCTTCCCACCACC	GGAGAAGTGTAGCTCCAGCC	ms10	7.2	1	[30]
SSR	AVRDC_MD997	CTGTTTGGAAGGAACCTTTA	TCAGTGATCCAATAATGTGC	ms10	20.8	1	[30]
CAPS	GMSK-CAPS	-	-	msk	0	-	[28]
dCAPS	SPGMS1	TGCAGAAATGGATC	AGGATTTACAAGGAGATGAA	ms_w	0	5	[15]
InDel	Indel-12	GCCTGAAGTTCAAGTGGCTC	CCACAGATGCAGTAACAAGCA	msc-1	0	2	[20]
dCAPS	dCAPs-msc-2	TTGTATCAATGGCTCATTGGAGATTGCGTCT	AACTCCCTCCCTGGCTACAAATATGTCC	msc-2	0	5	[21]
InDel	Ind198	TTCTCCTCCTGCTGTGCATG	TCTCCACCACAAGTCCCTCA	msc-3	0	10	[22]
SSR	H12	GTTCTATTAAGGGTGGTGAGGGT	CTCGCAGCAGATGGTAGTTTAGT	msc2	0.29	-	[32]
SSR	H24	TTAAACACGGGATGCAGCTACTA	AAGTTCCTTTTCGTGTAACGACC	msc2	0	-	[32]

标记类型	标记名称	CMS不育基因	引物序列		参考文献
标记类型	标记名称	CMS不育基因	前引物(5’-3’)	后引物(5’-3’)	参考文献
SCAR	atp6₆₀₇	atp6	AGTCCACTTGAACAATTTGAAATAATC	GTTCCGTACTTTACTTACGAGC	[38]
SCAR	coxII₇₀₈	coxII	GTCGGGAGAACTACCTAACTA	GGCTACCTAGTGATTTACAAGCA	[38]
SCAR	orf456-SCAR	orf456	ATGCCCAAAAGTCCCATGTA	TTACTCGGTTGCTCCATTGTTT	[36]
SCAR	SCAR₁₃₀	orf456	TTACGGCTCGTTACCGCAGC	AATTGACCGACCCGCCAT	[41]
-	accD-U	accD	GTGATTGGATTTCTATGCGG	GGAATGCCGTTTATTTGGCC	[39]
RT-PCR	orf300a	orf300a,orf314a	ATGTTCTCACTCTTAAAAAAATTAAGAGC	TTAGGATTTCCAATTCCAATGC	[42]

辣椒雄性不育分子研究进展

Molecular Advancements of Male Sterility in Pepper

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基因名称	定位区间/kb	定位区间长度/kb	两侧标记	候选基因	参考基因组	参考文献
Rf	无	821.00	4940-CAPS和PPR12-SCAR	CaPPR6	CM334 (V1.55)^a	[57]
Rf	214672-215170	498.60	pep20和pep43	无	Zunla-1 (V2.0)	[58]
Rf	214869-215727	858.26	P06-247和PW6-126 214	Capana06g002967和Capana06g002969	Zunla-1 (V2.0)	[59]
CaRf032	213924-214072	148.05	S1402和S1354	CA00g82510	CM334 (V1.55)	[46]
CaRfm	无	128.96	S409和S425	CaPPR6	CM334 (V1.55)	[47]
CaRf	217066-217281	270.10	CSSHjm9_44和CSSHjm9_60	Capana06g003028	Zunla-1 (V2.0)	[60]
Rf_u	4656-5054	398.00	G16-SCAR和13T7-SCAR	PPR	UCD10X (V1.0)	[61]
CaRf_HZ	215097-215631	533.81	P06gInDel-66和P06gInDel-89	Capana06g002968	Zunla-1 (V2.0)	[48]
Rf2	2561-2740	179.3	S1736 和 S1719	Capana06g000193	Zunla-1 (V2.0)	[45]
Rf	210577-215685	5100.00	KS18和KS22	Capana06g002866（NEDD8偶联酶基因）	Zunla-1 (V2.0)	[62]

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标记类型	标记名称	引物序列		基因	遗传距离/cM	参考文献
标记类型	标记名称	前引物(5’-3’)	后引物(5’-3’)	基因	遗传距离/cM	参考文献
RAPD	OP13₁₄₀₀	-	-	Rf	0.37	[50]
RAPD	OW19₈₀₀	-	-	Rf	8.12	[50]
CAPS	OPP13-CAPS	TACAGCTTCAAAGTAAACACAACC	ATTCGGGTCCCAAGAAGGTTCTAT	Rf	1.1	[51]
CAPS	AFRF8CAPS	GTTGATGCTCTATGGTTGGAGAAC	GCACTATTCCTATTGGCTTTCTG	Rf	1.8	[51]
SCAR	CRF3S1S	GTACACACCACTCGTCGCTCCT	TTCTTGGGTCCCTTTCTTCCAA	Rf	4.8/5.3	[52]
CAPS	PR-CAPS	ATGTCACCCCCACACACTCCTTCACCT	TCCCATCTAGCCTCTGCCTTCTCAAATG	Pr	1.8	[53]
SCAR	CRF-SCAR	GAAGTAGGCCAAAACTTATCACG	CACTAAGCCCGATGTATGAATC	Rf	1.4	[54]
STS	CaRf-FL-M2	ATGGATCCAAAGGACCACAAGCTAACC	GCATTGGGACTGAAATCCCCAAATAAG	Rf	0.5	[40]
CAPS	Co1Mod1-CAPS	CCACCCCAAACGTAAGGGATC	CAACTTAGCCAATACAATCCCAC	Rf	0	[57]

标记类型	标记名称	引物序列		基因	遗传距离/cM	参考文献
标记类型	标记名称	前引物(5’-3’)	后引物(5’-3’)	基因	遗传距离/cM	参考文献
RAPD	OP13₁₄₀₀	-	-	Rf	0.37	[50]
RAPD	OW19₈₀₀	-	-	Rf	8.12	[50]
CAPS	OPP13-CAPS	TACAGCTTCAAAGTAAACACAACC	ATTCGGGTCCCAAGAAGGTTCTAT	Rf	1.1	[51]
CAPS	AFRF8CAPS	GTTGATGCTCTATGGTTGGAGAAC	GCACTATTCCTATTGGCTTTCTG	Rf	1.8	[51]
SCAR	CRF3S1S	GTACACACCACTCGTCGCTCCT	TTCTTGGGTCCCTTTCTTCCAA	Rf	4.8/5.3	[52]
CAPS	PR-CAPS	ATGTCACCCCCACACACTCCTTCACCT	TCCCATCTAGCCTCTGCCTTCTCAAATG	Pr	1.8	[53]
SCAR	CRF-SCAR	GAAGTAGGCCAAAACTTATCACG	CACTAAGCCCGATGTATGAATC	Rf	1.4	[54]
STS	CaRf-FL-M2	ATGGATCCAAAGGACCACAAGCTAACC	GCATTGGGACTGAAATCCCCAAATAAG	Rf	0.5	[40]
CAPS	Co1Mod1-CAPS	CCACCCCAAACGTAAGGGATC	CAACTTAGCCAATACAATCCCAC	Rf	0	[57]