Research Progress of Tomato Male Sterility and Its Breeding Application

doi:10.16420/j.issn.0513-353x.2024-0759

Acta Horticulturae Sinica ›› 2025, Vol. 52 ›› Issue (8): 2081-2098.doi: 10.16420/j.issn.0513-353x.2024-0759

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Research Progress of Tomato Male Sterility and Its Breeding Application

ZHOU Ming¹, HAN Chenxu¹^,², YUAN Guoliang¹, FAN Xiaoqing¹, AI Pengfei²^,^*(), and LI Changbao¹^,^*()

¹ State Key Laboratory of Vegetable Biobreeding， National Engineering Research Center for Vegetables/Beijing Key Laboratory of Vegetable Germplasm Improvement/Key Laboratory of Biology and Genetic Improvement of Horticultural Crops（North China），Beijing Institute of Vegetable Science， Beijing Academy of Agriculture and Forestry Sciences Beijing 100097， China
² School of Food Science and Biology， Hebei University of Science and Technology， Shijiazhuang 050018， China

Received:2025-04-13 Revised:2025-06-22 Online:2025-08-25 Published:2025-08-19
Contact: AI Pengfei, and LI Changbao

Abstract

Abstract:

Tomato（Solanum lycopersicum）is one strictly self-pollinating crop，and its hybrid seed production requires artificial emasculation and pollination. With the increase in labor cost，male sterility based hybrid breeding system has become a promising technology. In recent years，with the advances in omics and molecular biotechnology，research on tomato male sterility has made rapid progress. This article reviewed the research progress on tomato male sterility and its breeding applications. The related studies on tomato male sterility mutants，male sterility genes，and male sterility regulation mechanisms were summarized. Moreover，the creation strategies and evaluation method of male sterile lines，male sterility propagation strategies，as well as elimination-free strategies were outlined. This article provided a reference for further research on tomato male sterility in the future.

Key words: tomato, male sterility, molecular mechanism, hybrid breeding application

ZHOU Ming, HAN Chenxu, YUAN Guoliang, FAN Xiaoqing, AI Pengfei, and LI Changbao. Research Progress of Tomato Male Sterility and Its Breeding Application[J]. Acta Horticulturae Sinica, 2025, 52(8): 2081-2098.

Figures/Tables 5

References 68

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类型Type	突变体 Mutant			表型Phenotype	参考文献Reference
结构不育 Structural male sterility	variable male sterile (vms)			高温雄性不育，低温部分恢复 Male sterility occurs under higher temperatures，and is restored partially under lower temperatures	Rick & Boynton,1967
	Sl-2/Slglo2/7b-1/tpi/ms30/ms33			雄蕊心皮化。sl 2高温不育，低温育性部分恢复。7b-1长日照下不育，短日照下育性部分恢复 Anthers develop abnormally with carpelloid-like stamens. sl 2 is male sterile under higher temperatures but show fertile under lower temperatures. 7b-1 shows male sterile in short days but fertile in long days.	Sawhney,1983；Pucci et al.,2017；Wei et al.,2024
	green pistillate (gpi)/ pi-2(pistillate 2)			雄蕊心皮化，同时伴有花瓣萼片化 Stamens are carpelloid，and petals are sepaloid	Rasmussen & Green,1993
	tag1			花药发育异常，雄蕊心皮化，花粉母细胞无法进行减数分裂，形成小孢子Anthers develop abnormally with carpelloid-like stamens，and pollen mother cells do not undergo meiosis to form microspores	Pnueli et al.,1994
	Slglo1/globosa/lepi/tpib			雄蕊发育异常。高温不育，低温育性部分恢复 Anthers develop abnormally. Male sterility occurs under higher temperature conditions，and fertility is restored under lower temperatures	Gómez et al.,1999
	sl/Sl-pr/ tap3/sldef/csp			雄蕊发育异常，高温不育，低温或喷施赤霉素育性部分恢复 Anthers develop abnormally. Male sterility occurs under higher temperature conditions，and fertility is restored partially under lower temperatures or with gibberellic acid treatment	Gómez et al.,1999；Li et al.,2024
结构不育 Structural male sterility	ms-15/tm6/tdr6		花药发育异常，雄蕊心皮化，花粉母细胞无法进行减数分裂，形成小孢子Anthers develop abnormally with carpelloid-like stamens，and pollen mother cells do not undergo meiosis to form microspores		Cao et al.,2019
功能不育 Functional male sterility	positional sterile (ps)		成熟花药无法开裂，花粉无法散出 The mature pollens are not released due to non-dehiscent anthers		Larson & Paur,1948
	exerted stigma (ex)		柱头外露 Exserted stigmas		Rick & Robinson,1951
	style2.1		柱头外露 Exserted stigmas		Chen et al.,2007
	ps2		药室外壁结构异常导致花药无法开裂，花粉无法散出 Locule wall of anthers develop abnormally，which lead to delayed anther dehiscence and failed pollen release		Gorguet et al.,2009
	procera(pro)		柱头外露exserted stigmas		Carrera et al.,2012
	Solanum lycopersicum long styles (SlLst)		柱头外露exserted stigmas		Cheng et al.,2021
孢子不育 Sporogenous male sterility	ms3		败育发生在减数分裂前。孢母细胞发育异常 Male sterility occurs before meiosis. Microspore mother cells develop abnormally		Rick & Butler,1956
	ms5		败育发生在减数分裂期。减数分裂异常，绒毡层细胞发育异常 Male sterility occurs during meiosis stage. Meiosis dysfunctions and tapetum develops abnormally		Rick & Butler,1956
	ms11		败育发生在减数分裂期。绒毡层提前降解，减数分裂异常，无法形成四分体 Male sterility occurs during meiosis stage. Tapetum degrades in advance. Microspore mother cells fail to give rise to tetrads owing to dysfunctional meiosis		Gorman et al.,1997
	Lesut2		败育发生时期不确定。花粉活力降低 The stage of male sterility occurrence is uncertain. The pollen viability reduces obviously		Hackel et al.,2005
	ms1		败育发生时期不确定。花粉完全败育。拟南芥、玉米、水稻ms1突变体绒毡层降解延迟，小孢子从四分体释放后花粉壁发育异常逐渐降解最终完全败育 The stage of male sterility occurrence is uncertain. The pollen is completely male sterile. The tapetums degradation is delayed and the microspores degenerate gradually owing to abnormal exine development，which lead to complete male sterility in ms1 of Arabidopsis，rice and corn.		Vizcay-Barrena & Wilson,2006；Yang et al.,2007,2019；An et al.,2020；Gökdemir et al.,2022
	ga20ox1		败育发生在减数分裂期。减数分裂异常，无法形成或只形成部分异常四分体，花粉部分败育。雌蕊发育异常 Male sterility occurs during meiosis stage. Owing to dysfunctional meiosis，microspore mother cells fail to produce tetrads or just produce few abnormal tetrads and generate few viable pollens. The mutant is also female sterile		Olimpieri et al.,2011
	ms10		败育发生在减数分裂期。减数分裂后期发生异常，绒毡层细胞质空泡化膨大，小孢子无法形成 Male sterility occurs during meiosis stage. Tapetum is enlarged and vacuolated，and meiocytes degrade and fail to give rise to microspores at anaphase		Jeong et al.,2014
	Lepro		败育发生时期不确定。花粉在柱头水合受到影响，花粉管伸长受抑制 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. The pollen hydration is affected，and the pollen tube elongation is inhibited		Yu & Mandayam,2014
	mapk7		败育发生在单核小孢子向二核小孢子转变期。绒毡层降解过快，花粉内壁发育异常，单核小孢子无法发育成成熟花粉，90%花粉无活力 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. The tapetum degrades in advance. Intine layer of pollen appears abnormal. And the uninucleate microspores can not develop into mature pollens. Approximately 90 % of the pollen grains lack viability		Chen et al.,2016
	gpat6-a		败育发生时期不确定。花粉数减少，花粉活力降低 The stage of male sterility occurrence is uncertain. The produced pollens decrease obviously，and pollen viability reduces.		Petit et al.,2016
	hyd		败育发生在减数分裂前。花药畸形，孢母细胞及绒毡层发育异常，无花粉产生 Male sterility occurs before meiosis. Microspore mother cells and tapetum layer develop abnormally in malformed anthers，and the mutant fail to produce any pollens		Rojas-Gracia et al.,2017
	mapk20		败育发生在单核小孢子向二核小孢子转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores		Chen et al.,2018
孢子不育 Sporogenous male sterility	Slmyb32	败育发生时期不确定。突变后花粉部分失活 The stage of male sterility occurrence is uncertain. The mutants can only produce partial inactivated pollens			Chen et al.,2018
孢子不育 Sporogenous male sterility	Slnced1	败育发生在减数分裂期。四分体发育异常，花粉部分败育。雌蕊发育异常 The mutants show male and female sterility. Male sterility occurs during meiosis stage. The mutants can only develop few normal pollens owing to abnormal tetrad development			Dai et al.,2018
	Slpin8	花药瘦小。败育发生时期不确定。80%花粉失去活力，花粉育性可部分被外源生长素恢复 The anthers are atrophied. Approximately 80% of the pollen grains are abnormal and lack viability，and the fertility is restored partially with IAA treatment			Gan et al.,2019
	ms32	花器变小。败育发生在减数分裂期。减数分裂异常，孢母细胞退化，无法形成四分体 The flower is smaller. Male sterility occurs during meiosis stage. Meiosis dysfunctions. Microspore mother cells degrade and can not generate tetrads			Liu et al.,2019
	Slida	败育发生时期不确定。绒毡层延迟降解，多数花粉失活，花药无法开裂 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. Most of pollens produced are inviable. The anthers are non-dehiscent			Wang et al.,2020
	bzr1	败育发生时期不确定。绒毡层降解延迟，花粉部分败育 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. The mutants can only produce partial inactivated pollens			Yan et al.,2020
	pif3	败育发生在单核小孢子向二核小孢子转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Yang et al.,2021
	Slglt1	败育发生在单核小孢子向二核小孢子转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Yang et al.,2021
	Slcwin9	败育发生在单核小孢子向二核小孢子转变期花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Yang et al.,2021
	ams	败育发生时期不确定。花粉发育异常，多数花粉失去活性 The stage of male sterility occurrence is uncertain. The pollens develop abnormally and most of them are inviable			Bao et al.,2022
	lerboh lerbohe	败育发生时期不确定。绒毡层降解延迟，花粉败育 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. The mutants can only produce partial inactivated pollens			Dai et al.,2022
	tpd1	败育发生在减数分裂前。绒毡层无法形成，孢母细胞无法进行减数分裂 Male sterility occurs before meiosis. Microspore mother cells fail to undergo meiosis and tapetum degrade grandually			Salazar-Sarasua et al.,2022
	Slaco2	败育发生在减数分裂期。花粉形态结构正常，但活力极低 Male sterility occurs during meiosis stage. The mutants produce normal-shaped pollens，but the pollen viability is extremely low			Secgin et al.,2022
	mapk4	败育发生在二核小孢子向成熟花粉转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Wang et al.,2022
	bg10	败育发生在四分体向小孢子转变期。胼胝质降解受阻，小孢子无法从四分体解离，花粉畸形，活力和发芽率极低 Male sterility occurs during the transition stage from tetrads to microspores. Callose degradation is inhibited，and microspores can not be released from tetrads. The pollens are abnormal and almost inviable			Pei et al.,2023

类型Type	突变体 Mutant			表型Phenotype	参考文献Reference
结构不育 Structural male sterility	variable male sterile (vms)			高温雄性不育，低温部分恢复 Male sterility occurs under higher temperatures，and is restored partially under lower temperatures	Rick & Boynton,1967
	Sl-2/Slglo2/7b-1/tpi/ms30/ms33			雄蕊心皮化。sl 2高温不育，低温育性部分恢复。7b-1长日照下不育，短日照下育性部分恢复 Anthers develop abnormally with carpelloid-like stamens. sl 2 is male sterile under higher temperatures but show fertile under lower temperatures. 7b-1 shows male sterile in short days but fertile in long days.	Sawhney,1983；Pucci et al.,2017；Wei et al.,2024
	green pistillate (gpi)/ pi-2(pistillate 2)			雄蕊心皮化，同时伴有花瓣萼片化 Stamens are carpelloid，and petals are sepaloid	Rasmussen & Green,1993
	tag1			花药发育异常，雄蕊心皮化，花粉母细胞无法进行减数分裂，形成小孢子Anthers develop abnormally with carpelloid-like stamens，and pollen mother cells do not undergo meiosis to form microspores	Pnueli et al.,1994
	Slglo1/globosa/lepi/tpib			雄蕊发育异常。高温不育，低温育性部分恢复 Anthers develop abnormally. Male sterility occurs under higher temperature conditions，and fertility is restored under lower temperatures	Gómez et al.,1999
	sl/Sl-pr/ tap3/sldef/csp			雄蕊发育异常，高温不育，低温或喷施赤霉素育性部分恢复 Anthers develop abnormally. Male sterility occurs under higher temperature conditions，and fertility is restored partially under lower temperatures or with gibberellic acid treatment	Gómez et al.,1999；Li et al.,2024
结构不育 Structural male sterility	ms-15/tm6/tdr6		花药发育异常，雄蕊心皮化，花粉母细胞无法进行减数分裂，形成小孢子Anthers develop abnormally with carpelloid-like stamens，and pollen mother cells do not undergo meiosis to form microspores		Cao et al.,2019
功能不育 Functional male sterility	positional sterile (ps)		成熟花药无法开裂，花粉无法散出 The mature pollens are not released due to non-dehiscent anthers		Larson & Paur,1948
	exerted stigma (ex)		柱头外露 Exserted stigmas		Rick & Robinson,1951
	style2.1		柱头外露 Exserted stigmas		Chen et al.,2007
	ps2		药室外壁结构异常导致花药无法开裂，花粉无法散出 Locule wall of anthers develop abnormally，which lead to delayed anther dehiscence and failed pollen release		Gorguet et al.,2009
	procera(pro)		柱头外露exserted stigmas		Carrera et al.,2012
	Solanum lycopersicum long styles (SlLst)		柱头外露exserted stigmas		Cheng et al.,2021
孢子不育 Sporogenous male sterility	ms3		败育发生在减数分裂前。孢母细胞发育异常 Male sterility occurs before meiosis. Microspore mother cells develop abnormally		Rick & Butler,1956
	ms5		败育发生在减数分裂期。减数分裂异常，绒毡层细胞发育异常 Male sterility occurs during meiosis stage. Meiosis dysfunctions and tapetum develops abnormally		Rick & Butler,1956
	ms11		败育发生在减数分裂期。绒毡层提前降解，减数分裂异常，无法形成四分体 Male sterility occurs during meiosis stage. Tapetum degrades in advance. Microspore mother cells fail to give rise to tetrads owing to dysfunctional meiosis		Gorman et al.,1997
	Lesut2		败育发生时期不确定。花粉活力降低 The stage of male sterility occurrence is uncertain. The pollen viability reduces obviously		Hackel et al.,2005
	ms1		败育发生时期不确定。花粉完全败育。拟南芥、玉米、水稻ms1突变体绒毡层降解延迟，小孢子从四分体释放后花粉壁发育异常逐渐降解最终完全败育 The stage of male sterility occurrence is uncertain. The pollen is completely male sterile. The tapetums degradation is delayed and the microspores degenerate gradually owing to abnormal exine development，which lead to complete male sterility in ms1 of Arabidopsis，rice and corn.		Vizcay-Barrena & Wilson,2006；Yang et al.,2007,2019；An et al.,2020；Gökdemir et al.,2022
	ga20ox1		败育发生在减数分裂期。减数分裂异常，无法形成或只形成部分异常四分体，花粉部分败育。雌蕊发育异常 Male sterility occurs during meiosis stage. Owing to dysfunctional meiosis，microspore mother cells fail to produce tetrads or just produce few abnormal tetrads and generate few viable pollens. The mutant is also female sterile		Olimpieri et al.,2011
	ms10		败育发生在减数分裂期。减数分裂后期发生异常，绒毡层细胞质空泡化膨大，小孢子无法形成 Male sterility occurs during meiosis stage. Tapetum is enlarged and vacuolated，and meiocytes degrade and fail to give rise to microspores at anaphase		Jeong et al.,2014
	Lepro		败育发生时期不确定。花粉在柱头水合受到影响，花粉管伸长受抑制 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. The pollen hydration is affected，and the pollen tube elongation is inhibited		Yu & Mandayam,2014
	mapk7		败育发生在单核小孢子向二核小孢子转变期。绒毡层降解过快，花粉内壁发育异常，单核小孢子无法发育成成熟花粉，90%花粉无活力 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. The tapetum degrades in advance. Intine layer of pollen appears abnormal. And the uninucleate microspores can not develop into mature pollens. Approximately 90 % of the pollen grains lack viability		Chen et al.,2016
	gpat6-a		败育发生时期不确定。花粉数减少，花粉活力降低 The stage of male sterility occurrence is uncertain. The produced pollens decrease obviously，and pollen viability reduces.		Petit et al.,2016
	hyd		败育发生在减数分裂前。花药畸形，孢母细胞及绒毡层发育异常，无花粉产生 Male sterility occurs before meiosis. Microspore mother cells and tapetum layer develop abnormally in malformed anthers，and the mutant fail to produce any pollens		Rojas-Gracia et al.,2017
	mapk20		败育发生在单核小孢子向二核小孢子转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores		Chen et al.,2018
孢子不育 Sporogenous male sterility	Slmyb32	败育发生时期不确定。突变后花粉部分失活 The stage of male sterility occurrence is uncertain. The mutants can only produce partial inactivated pollens			Chen et al.,2018
孢子不育 Sporogenous male sterility	Slnced1	败育发生在减数分裂期。四分体发育异常，花粉部分败育。雌蕊发育异常 The mutants show male and female sterility. Male sterility occurs during meiosis stage. The mutants can only develop few normal pollens owing to abnormal tetrad development			Dai et al.,2018
	Slpin8	花药瘦小。败育发生时期不确定。80%花粉失去活力，花粉育性可部分被外源生长素恢复 The anthers are atrophied. Approximately 80% of the pollen grains are abnormal and lack viability，and the fertility is restored partially with IAA treatment			Gan et al.,2019
	ms32	花器变小。败育发生在减数分裂期。减数分裂异常，孢母细胞退化，无法形成四分体 The flower is smaller. Male sterility occurs during meiosis stage. Meiosis dysfunctions. Microspore mother cells degrade and can not generate tetrads			Liu et al.,2019
	Slida	败育发生时期不确定。绒毡层延迟降解，多数花粉失活，花药无法开裂 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. Most of pollens produced are inviable. The anthers are non-dehiscent			Wang et al.,2020
	bzr1	败育发生时期不确定。绒毡层降解延迟，花粉部分败育 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. The mutants can only produce partial inactivated pollens			Yan et al.,2020
	pif3	败育发生在单核小孢子向二核小孢子转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Yang et al.,2021
	Slglt1	败育发生在单核小孢子向二核小孢子转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Yang et al.,2021
	Slcwin9	败育发生在单核小孢子向二核小孢子转变期花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Yang et al.,2021
	ams	败育发生时期不确定。花粉发育异常，多数花粉失去活性 The stage of male sterility occurrence is uncertain. The pollens develop abnormally and most of them are inviable			Bao et al.,2022
	lerboh lerbohe	败育发生时期不确定。绒毡层降解延迟，花粉败育 The stage of male sterility occurrence is uncertain. The tapetums degradation is delayed. The mutants can only produce partial inactivated pollens			Dai et al.,2022
	tpd1	败育发生在减数分裂前。绒毡层无法形成，孢母细胞无法进行减数分裂 Male sterility occurs before meiosis. Microspore mother cells fail to undergo meiosis and tapetum degrade grandually			Salazar-Sarasua et al.,2022
	Slaco2	败育发生在减数分裂期。花粉形态结构正常，但活力极低 Male sterility occurs during meiosis stage. The mutants produce normal-shaped pollens，but the pollen viability is extremely low			Secgin et al.,2022
	mapk4	败育发生在二核小孢子向成熟花粉转变期。花粉由单核向二核转变时发生败育 Male sterility occurs during the transition stage from uninucleate to binucleate microspores. Uninucleate microspores fail to undergo mitosis to generate binucleate microspores			Wang et al.,2022
	bg10	败育发生在四分体向小孢子转变期。胼胝质降解受阻，小孢子无法从四分体解离，花粉畸形，活力和发芽率极低 Male sterility occurs during the transition stage from tetrads to microspores. Callose degradation is inhibited，and microspores can not be released from tetrads. The pollens are abnormal and almost inviable			Pei et al.,2023

类型 Type	基因/位点 Gene/locus	基因编号 ID	编码蛋白 Protein	表达 Expression	调控机理 Molecular regulatory mechanism	参考文献 Reference
结构不育 Structural male sterility	TAP3/SlDEF	Solyc04g081000	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达 Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Quinet et al.,2014；Li et al.,2024
	SlGLO1/GLOBOSA,/LePI/TPIB	Solyc08g06723	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Guo et al.,2016
	7B-1/GLO2/MS30/MS33	Solyc06g059970	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Pucci et al.,2017；Wei et al.,2024
	MS-15/TM6/TDR6	Solyc02g084630	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达 Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Cao et al.,2019
功能不育 Functional male sterility	Style2.1	Solyc02g087660	Style2.1，basic helix-loop-belix (bHLH)转录因子 bHLH family transcription factor	在花柱组织细胞内表达 Expressed in style through the whole development satge	调控花柱细胞伸长，柱头外露 Controlling style cell elongation to protrude the style beyond the anthers	Chen et al.,2007
	PS-2	Solyc04g015530	多聚半乳糖醛酸酶 Polygalacturonase	拟南芥同源基因在花药开裂区域表达Arabidopsis homolog of PS-2 is expressed in the anther dehiscence zone	未知 Unknown	Gorguet et al.,2009
孢子不育 Sporogenous male sterility	LeSUT2	Solyc11g017010	蔗糖转运蛋白2 Sucrose transporter2	花药组织内高表达 Highly expressed in stamens	可能具有蔗糖转运蛋白活性，参与蔗糖运输Probably participate in sucrose transport as the transporter protein	Hackel et al 2005
	MS1	Solyc04g008420	Male sterility1，plant homeodomain（PHD）转录因子 PHD transcription factor	拟南芥同源基因在减数分裂期绒毡层表达 Arabidopsis homolog of MS1 is expressed in tapetum	拟南芥和水稻同源基因调控花药脂质代谢，调控花粉壁发育相关的基因表达 Arabidopsis and rice homologs of MS1 regulate transcription of genes involved in lipid metabolism，programmed cell death of tapetum and pollen wall formation	Yang et al.,2007；Gökdemir et al.,2022； Yang et al.,2019
	GA20ox1	Solyc03g006880	GA 20 氧化酶1，参与 GA 代谢 Gibberellins 20 oxidase1，involved in Gibberellins metabolism	花药组织内高表达 Highly expressed in stamens	赤霉素信号转导受阻导致减数分裂异常 Inhibition of GA signaling leads to abnormal meiosis	Olimpieri et al.,2011
	SlDELLA	Solyc11g011260	DELLA蛋白，GA信号转导元件 GA signal transduction element	未知 Unknown	通过赤霉素信号转导调控花柱细胞伸长，柱头外露 Controlling style cell elongation to protrude the style beyond the anthers through GA signaling transduction	Carrera et al.,2012
	MS10	Solyc02g079810	Male sterility10，basic helix-loop-helix (bHLH) 家族转录因子 bHLH family transcription factor	减数分裂以前在花粉母细胞、绒毡层细胞内表达；减数分裂期主要在绒毡层细胞内表达 Expressed in microspore mother cells and tapetums before meiosis. Expressed in tapetums during meiosis stage	调控参与花粉发育相关的细胞壁形成、脂类代谢基因以及减数分裂相关基因 Regulating expression of genes involved in meiosis，pollen wall formation and lipid metabolism	Jeong et al.,2014
	LePro1	Solyc12g044630	花粉肌动蛋白1 Pollen Profilin 1	在成熟花粉或花粉管中表达Expressed in mature pollen and pollen tubes	调控花粉在柱头的水合、萌发以及花粉管生长，花粉活力降低 Controlling pollen hydration，germination and pollen tube elongation	Yu & Mandayam,2014
	SlMAPK7	Solyc08g081490	裂原活化蛋白激酶7 Mitogen-activated protein kinase 7	未知 Unknown	调控绒毡层程序化死亡和花粉内壁发育	Chen et al.,2016
	Sl-IAA27	Solyc03g120500	Aux/IAA27 蛋白生长素信号转导元件Aux/IAA蛋白 auxin (IAA) signal transduction element	未知 Unknown	生长素信号转导受阻导致减数分裂异常 Auxin (IAA) signaling is blocked in the mutant，which leads to defect meiosis	Guillotin et al.,2016
	GPAT6-a	Solyc09g014350	甘油-3-磷酸酰基转移酶 Glycerol-3-phosphate acyltransferase	未知 Unknown	可能参与花粉发育过程中脂质、细胞壁生物合成及代谢 Probably regulating pollen wall formation and lipid metabolism	Petit et al.,2016
	HYD	Solyc07g063670	HYDRA，SPOROCYTELESS/NOZZLE (SPL/NZZ) 转录因子 SPOROCYTELESS/NOZZLE (SPL/NZZ) transcription factor	发育早期在雄蕊组织各部位表达，之后在孢母细胞内表达 Expressed in whole stamen tissues at early stage，then in microspore mother cells	可能通过调控赤霉素合成影响雄蕊发育 Regulating stamen development through the modulation of gibberellin metabolism	Rojas-Gracia et al.,2017
	SlMAPK20	Solyc07g056350	裂原活化蛋白激酶 Mitogen-activated protein kinase20	主要在四分体以及小孢子细胞中表达 Expressed in tetrads and microspores	与SlMYB32协同转录调控蔗糖、生长素代谢和其它信号途径基因表达，调控减数分裂后花粉发育 Interacting with SlMYB32 to regulate transcription of genes involved in sucrose and IAA metabolism，as well as other signaling pathway after meiosis	Chen et al.,2018
	SlMYB32	Solyc10g055410	MYB32，R2R3 MYB 转录因子 R2R3MYB transcription factor 32	二核花粉时期在花药组织内高表达 Expressed in stamens at binucleate microspores stage	未知 Unknown	Chen et al.,2018
	SlNCED1	Solyc08g016720	9-顺式环氧类胡萝卜素双加氧酶1，参与ABA 合成 9-cis-epoxycarotenoid dioxygenase1，involved in ABA synthesis	转录发生在花粉发育整个时期生殖细胞内 Expressed in germ cells through the whole pollen development stage	ABA信号调控花粉发育相关的脂类、碳水化合物以及细胞壁形成基因表达 Regulating expression of genes involved in lipid metabolism，carbohydrate metabolism and pollen wall formation through ABA signaling	Dai et al.,2018
	SlPIN8	Solyc02g087660	PIN-FORMED 8 蛋白，生长素转运蛋白 auxin (IAA) transport protein	花粉发育后期在花粉细胞内表达 Expressed in germ cells at late-stage pollen development	调控生长素稳态；正调控MS10基因表达。Regulating cellular auxin homoeostasis；Positively regulating MS10 gene expression	Gan et al.,2019
	MS32	Solyc01g081100	Male sterility1，basic helix-loop-helix (bHLH) 家族转录因子 bHLH family transcription factor	未知 Unknown	拟南芥同源基因DYT1-TDF1- AMS-bHLH10/89/ 91-MYB80 形成转录模块调控花粉和绒毡层发育 Arabidopsis homologs of DYT1-TDF1-AMS-bHLH10 /89/ 91-MYB80 form a transcriptional cascade to regulate pollen and tapetum development	Liu et al.,2019
	STR	Solyc03g053130	异胡豆苷合酶 Strictosidine synthase	未知 Unknown	拟南芥同源基因为花粉外壁发育代谢通路的关键基因 Arabidopsis homolog of STR is the key gene in metabolic pathway of pollen wall development	Du et al.,2020
	SlIDA	Solyc05g010000	14-mer烯醇化酶纤溶酶原互作多肽 14-mer enolase- plasminogen interaction peptide peptide	发育前期在绒毡层、小孢子表达；发育后期在花药隔壁和成熟花粉以及花粉管内表达 Expressed in the tapetum and microspores during early stage；Expressed in septum，mature pollens and pollen tube during late stage	正调控花粉及绒毡层内的活性氧水平 Positively regulating ROS levels in tapetums and pollens	Wang et al.,2020
	BZR1	Solyc04g079980	BRASSINAZOLE RESISTANT 1，油菜素内酯信号转录因子 Brassinosteroid (BR) signaling transcription factor	未知 Unknown	影响了绒毡层活性氧和程序化死亡，导致绒毡层降解延迟或提早 Altering the timing of reactive oxygen species (ROS) production and programmed cell death (PCD) in tapetal cells，resulting in delayed or premature tapetal degeneration	Yan et al.,2020
	DWF	Solyc02g089160	CYP85A1，细胞色素P450 Cytochrome P450	未知 Unknown	影响了绒毡层活性氧和程序化死亡，导致绒毡层降解延迟或提早 Altering the timing of reactive oxygen species (ROS) production and programmed cell death (PCD) in tapetal cells，resulting in delayed or premature tapetal degeneration	Yan et al.,2020
	PIF3	Solyc01g102300	光敏色素互作因子3，basic helix-loop-helix (bHLH) 家族转录因子 Phytochrome interacting factor3，bHLH family transcription factor	转录发生在花粉发育整个时期生殖细胞内 Expressed in germ cells through the whole pollen development stage	转录调控花粉发育相关的碳水化合物代谢以及生长素信号途径基因 Regulating transcription of genes involved in carbohydrate metabolism and IAA signaling	Yang et al.,2021
	SlGLT1	Solyc03g083440	NADH依赖的谷氨酰胺-氧戊二酸转氨酶 NADH-dependent glutamine-2-oxoglutarate aminotransferase	主要在四分体及成熟花粉 Expressed in tetrads and mature pollens	可能参与生长素合成前体代谢 Probably involved in synthesis of auxin precursors	Yang et al.,2021
	SlCWIN9	Solyc10g085650	细胞壁酸性转化酶9 Cell wall invertase 9	主要在四分体及成熟花粉表达 Expressed in tetrads and mature pollens	可能参与花粉发育相关的糖代谢 Regulating sugar metabolism involved in pollen development	Yang et al.,2021
	SlAMS	Solyc08g062780	ABORTED MICROSPORES，bHLH 转录因子 bHLH family transcription factor	花药组织内表达 Expressed specially in stamens	拟南芥同源基因DYT1-TDF1- AMS-bHLH10/89/ 91-MYB80形成花粉、绒毡层发育转录调控模块 Arabidopsis homologs of DYT1-TDF1-AMS- bHLH10/89/ 91-MYB80 form a transcriptional cascade to regulate pollen and tapetum development	Bao et al.,2022
	LeRBOHE	Solyc07g042460	呼吸爆发氧化酶Respiratory burst oxidase	花药组织内表达 Highly expressed in stamens	正调控花药组织内活性氧水平，负调控绒毡层程序化死亡 Regulating ROS levels in anthers positively and regulating programmed cell death of tapetum negatively	Dai et al.,2022
	SlCMT4	Solyc08g005400	DNA甲基转移酶4 DNA methyltransferase 4	未知 Unknown	转录调控花粉壁形成相关的果胶甲基酯酶抑制蛋白基因SlPMEI 和花粉管发育相关的前蛋白质原基因SlPRALF的表达，控制花粉活力和花粉管伸长 Transcriptionally regulating the expression of the pectin methylesterase inhibitor gene SlPMEI involved in pollen wall formation and the preproprotein gene SlPRALF that negatively regulates pollen tube elongation	Guo et al.,2022
	SlTPD1	Solyc03g097530	Tapetum determinant1,小肽，拟南芥同源蛋白为EMS1/EXS 配体 Tapetum determinant1，small peptide，Arabidopsis homolog of TPD1 acts as a ligand of EXCESS MICROSPOROCYTES 1/EXTRA SPOROGENOUS CELLS (EMS1/EXS)	转录发生在发育各时期，主要在孢原细胞、绒毡层组织、花粉细胞表达 Expressed in sporogenous cells，tapetum，microspores and mature pollens of anthers at different stages of pollen development	调控花药组织内活性氧稳态，调控花粉发育相关的多个基因如MS10，AMS等 Regulating redox homeostasis during male gametogenesis；Regulating transcription of genes involved in pollen development，such as MS10，AMS and so on	Salazar-Sarasua et al.,2022
	SlACO2	Solyc12g005860	ACC合成酶 1-aminocyclopropane-1- carboxylicacidoxidase，ACO	未知 Unknown	可能通过乙烯信号调控控制花粉发育 Probably regulating pollen development through ethylene signaling	Secgin et al.,2022
	SlMAPK4	Solyc11g072630	裂原活化蛋白激酶 Mitogen-activated protein kinase 4	仅在成熟花粉粒中表达 Expressed in mature pollens	调控花粉发育相关的碳水化合物代谢基因、信号转导基因。调控花粉壁发育 Involved in pollen wall development. Regulating gens of carbohydrate metabolism and signaling transduction	Wang et al.,2022
	SlBG10	Solyc11g065280	β-1,3-葡聚糖酶β-1,3-glucanase	发育早期在幼蕾期组织内高表达Highly expressed in young stamens at early development stage	具有胼胝质水解酶活性，参与胼胝质代谢 Involved in callus metabolism as callus hydrolase	Pei et al.,2023

类型 Type	基因/位点 Gene/locus	基因编号 ID	编码蛋白 Protein	表达 Expression	调控机理 Molecular regulatory mechanism	参考文献 Reference
结构不育 Structural male sterility	TAP3/SlDEF	Solyc04g081000	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达 Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Quinet et al.,2014；Li et al.,2024
	SlGLO1/GLOBOSA,/LePI/TPIB	Solyc08g06723	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Guo et al.,2016
	7B-1/GLO2/MS30/MS33	Solyc06g059970	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Pucci et al.,2017；Wei et al.,2024
	MS-15/TM6/TDR6	Solyc02g084630	MADS-box家族B类转录因子 Class B MADS-box transcription factors	发育初期在花药组织内表达 Expressed in early stage of stamen development	调控花器官发育的ABCDE分子模型 ABCDE model of flower development	Cao et al.,2019
功能不育 Functional male sterility	Style2.1	Solyc02g087660	Style2.1，basic helix-loop-belix (bHLH)转录因子 bHLH family transcription factor	在花柱组织细胞内表达 Expressed in style through the whole development satge	调控花柱细胞伸长，柱头外露 Controlling style cell elongation to protrude the style beyond the anthers	Chen et al.,2007
	PS-2	Solyc04g015530	多聚半乳糖醛酸酶 Polygalacturonase	拟南芥同源基因在花药开裂区域表达Arabidopsis homolog of PS-2 is expressed in the anther dehiscence zone	未知 Unknown	Gorguet et al.,2009
孢子不育 Sporogenous male sterility	LeSUT2	Solyc11g017010	蔗糖转运蛋白2 Sucrose transporter2	花药组织内高表达 Highly expressed in stamens	可能具有蔗糖转运蛋白活性，参与蔗糖运输Probably participate in sucrose transport as the transporter protein	Hackel et al 2005
	MS1	Solyc04g008420	Male sterility1，plant homeodomain（PHD）转录因子 PHD transcription factor	拟南芥同源基因在减数分裂期绒毡层表达 Arabidopsis homolog of MS1 is expressed in tapetum	拟南芥和水稻同源基因调控花药脂质代谢，调控花粉壁发育相关的基因表达 Arabidopsis and rice homologs of MS1 regulate transcription of genes involved in lipid metabolism，programmed cell death of tapetum and pollen wall formation	Yang et al.,2007；Gökdemir et al.,2022； Yang et al.,2019
	GA20ox1	Solyc03g006880	GA 20 氧化酶1，参与 GA 代谢 Gibberellins 20 oxidase1，involved in Gibberellins metabolism	花药组织内高表达 Highly expressed in stamens	赤霉素信号转导受阻导致减数分裂异常 Inhibition of GA signaling leads to abnormal meiosis	Olimpieri et al.,2011
	SlDELLA	Solyc11g011260	DELLA蛋白，GA信号转导元件 GA signal transduction element	未知 Unknown	通过赤霉素信号转导调控花柱细胞伸长，柱头外露 Controlling style cell elongation to protrude the style beyond the anthers through GA signaling transduction	Carrera et al.,2012
	MS10	Solyc02g079810	Male sterility10，basic helix-loop-helix (bHLH) 家族转录因子 bHLH family transcription factor	减数分裂以前在花粉母细胞、绒毡层细胞内表达；减数分裂期主要在绒毡层细胞内表达 Expressed in microspore mother cells and tapetums before meiosis. Expressed in tapetums during meiosis stage	调控参与花粉发育相关的细胞壁形成、脂类代谢基因以及减数分裂相关基因 Regulating expression of genes involved in meiosis，pollen wall formation and lipid metabolism	Jeong et al.,2014
	LePro1	Solyc12g044630	花粉肌动蛋白1 Pollen Profilin 1	在成熟花粉或花粉管中表达Expressed in mature pollen and pollen tubes	调控花粉在柱头的水合、萌发以及花粉管生长，花粉活力降低 Controlling pollen hydration，germination and pollen tube elongation	Yu & Mandayam,2014
	SlMAPK7	Solyc08g081490	裂原活化蛋白激酶7 Mitogen-activated protein kinase 7	未知 Unknown	调控绒毡层程序化死亡和花粉内壁发育	Chen et al.,2016
	Sl-IAA27	Solyc03g120500	Aux/IAA27 蛋白生长素信号转导元件Aux/IAA蛋白 auxin (IAA) signal transduction element	未知 Unknown	生长素信号转导受阻导致减数分裂异常 Auxin (IAA) signaling is blocked in the mutant，which leads to defect meiosis	Guillotin et al.,2016
	GPAT6-a	Solyc09g014350	甘油-3-磷酸酰基转移酶 Glycerol-3-phosphate acyltransferase	未知 Unknown	可能参与花粉发育过程中脂质、细胞壁生物合成及代谢 Probably regulating pollen wall formation and lipid metabolism	Petit et al.,2016
	HYD	Solyc07g063670	HYDRA，SPOROCYTELESS/NOZZLE (SPL/NZZ) 转录因子 SPOROCYTELESS/NOZZLE (SPL/NZZ) transcription factor	发育早期在雄蕊组织各部位表达，之后在孢母细胞内表达 Expressed in whole stamen tissues at early stage，then in microspore mother cells	可能通过调控赤霉素合成影响雄蕊发育 Regulating stamen development through the modulation of gibberellin metabolism	Rojas-Gracia et al.,2017
	SlMAPK20	Solyc07g056350	裂原活化蛋白激酶 Mitogen-activated protein kinase20	主要在四分体以及小孢子细胞中表达 Expressed in tetrads and microspores	与SlMYB32协同转录调控蔗糖、生长素代谢和其它信号途径基因表达，调控减数分裂后花粉发育 Interacting with SlMYB32 to regulate transcription of genes involved in sucrose and IAA metabolism，as well as other signaling pathway after meiosis	Chen et al.,2018
	SlMYB32	Solyc10g055410	MYB32，R2R3 MYB 转录因子 R2R3MYB transcription factor 32	二核花粉时期在花药组织内高表达 Expressed in stamens at binucleate microspores stage	未知 Unknown	Chen et al.,2018
	SlNCED1	Solyc08g016720	9-顺式环氧类胡萝卜素双加氧酶1，参与ABA 合成 9-cis-epoxycarotenoid dioxygenase1，involved in ABA synthesis	转录发生在花粉发育整个时期生殖细胞内 Expressed in germ cells through the whole pollen development stage	ABA信号调控花粉发育相关的脂类、碳水化合物以及细胞壁形成基因表达 Regulating expression of genes involved in lipid metabolism，carbohydrate metabolism and pollen wall formation through ABA signaling	Dai et al.,2018
	SlPIN8	Solyc02g087660	PIN-FORMED 8 蛋白，生长素转运蛋白 auxin (IAA) transport protein	花粉发育后期在花粉细胞内表达 Expressed in germ cells at late-stage pollen development	调控生长素稳态；正调控MS10基因表达。Regulating cellular auxin homoeostasis；Positively regulating MS10 gene expression	Gan et al.,2019
	MS32	Solyc01g081100	Male sterility1，basic helix-loop-helix (bHLH) 家族转录因子 bHLH family transcription factor	未知 Unknown	拟南芥同源基因DYT1-TDF1- AMS-bHLH10/89/ 91-MYB80 形成转录模块调控花粉和绒毡层发育 Arabidopsis homologs of DYT1-TDF1-AMS-bHLH10 /89/ 91-MYB80 form a transcriptional cascade to regulate pollen and tapetum development	Liu et al.,2019
	STR	Solyc03g053130	异胡豆苷合酶 Strictosidine synthase	未知 Unknown	拟南芥同源基因为花粉外壁发育代谢通路的关键基因 Arabidopsis homolog of STR is the key gene in metabolic pathway of pollen wall development	Du et al.,2020
	SlIDA	Solyc05g010000	14-mer烯醇化酶纤溶酶原互作多肽 14-mer enolase- plasminogen interaction peptide peptide	发育前期在绒毡层、小孢子表达；发育后期在花药隔壁和成熟花粉以及花粉管内表达 Expressed in the tapetum and microspores during early stage；Expressed in septum，mature pollens and pollen tube during late stage	正调控花粉及绒毡层内的活性氧水平 Positively regulating ROS levels in tapetums and pollens	Wang et al.,2020
	BZR1	Solyc04g079980	BRASSINAZOLE RESISTANT 1，油菜素内酯信号转录因子 Brassinosteroid (BR) signaling transcription factor	未知 Unknown	影响了绒毡层活性氧和程序化死亡，导致绒毡层降解延迟或提早 Altering the timing of reactive oxygen species (ROS) production and programmed cell death (PCD) in tapetal cells，resulting in delayed or premature tapetal degeneration	Yan et al.,2020
	DWF	Solyc02g089160	CYP85A1，细胞色素P450 Cytochrome P450	未知 Unknown	影响了绒毡层活性氧和程序化死亡，导致绒毡层降解延迟或提早 Altering the timing of reactive oxygen species (ROS) production and programmed cell death (PCD) in tapetal cells，resulting in delayed or premature tapetal degeneration	Yan et al.,2020
	PIF3	Solyc01g102300	光敏色素互作因子3，basic helix-loop-helix (bHLH) 家族转录因子 Phytochrome interacting factor3，bHLH family transcription factor	转录发生在花粉发育整个时期生殖细胞内 Expressed in germ cells through the whole pollen development stage	转录调控花粉发育相关的碳水化合物代谢以及生长素信号途径基因 Regulating transcription of genes involved in carbohydrate metabolism and IAA signaling	Yang et al.,2021
	SlGLT1	Solyc03g083440	NADH依赖的谷氨酰胺-氧戊二酸转氨酶 NADH-dependent glutamine-2-oxoglutarate aminotransferase	主要在四分体及成熟花粉 Expressed in tetrads and mature pollens	可能参与生长素合成前体代谢 Probably involved in synthesis of auxin precursors	Yang et al.,2021
	SlCWIN9	Solyc10g085650	细胞壁酸性转化酶9 Cell wall invertase 9	主要在四分体及成熟花粉表达 Expressed in tetrads and mature pollens	可能参与花粉发育相关的糖代谢 Regulating sugar metabolism involved in pollen development	Yang et al.,2021
	SlAMS	Solyc08g062780	ABORTED MICROSPORES，bHLH 转录因子 bHLH family transcription factor	花药组织内表达 Expressed specially in stamens	拟南芥同源基因DYT1-TDF1- AMS-bHLH10/89/ 91-MYB80形成花粉、绒毡层发育转录调控模块 Arabidopsis homologs of DYT1-TDF1-AMS- bHLH10/89/ 91-MYB80 form a transcriptional cascade to regulate pollen and tapetum development	Bao et al.,2022
	LeRBOHE	Solyc07g042460	呼吸爆发氧化酶Respiratory burst oxidase	花药组织内表达 Highly expressed in stamens	正调控花药组织内活性氧水平，负调控绒毡层程序化死亡 Regulating ROS levels in anthers positively and regulating programmed cell death of tapetum negatively	Dai et al.,2022
	SlCMT4	Solyc08g005400	DNA甲基转移酶4 DNA methyltransferase 4	未知 Unknown	转录调控花粉壁形成相关的果胶甲基酯酶抑制蛋白基因SlPMEI 和花粉管发育相关的前蛋白质原基因SlPRALF的表达，控制花粉活力和花粉管伸长 Transcriptionally regulating the expression of the pectin methylesterase inhibitor gene SlPMEI involved in pollen wall formation and the preproprotein gene SlPRALF that negatively regulates pollen tube elongation	Guo et al.,2022
	SlTPD1	Solyc03g097530	Tapetum determinant1,小肽，拟南芥同源蛋白为EMS1/EXS 配体 Tapetum determinant1，small peptide，Arabidopsis homolog of TPD1 acts as a ligand of EXCESS MICROSPOROCYTES 1/EXTRA SPOROGENOUS CELLS (EMS1/EXS)	转录发生在发育各时期，主要在孢原细胞、绒毡层组织、花粉细胞表达 Expressed in sporogenous cells，tapetum，microspores and mature pollens of anthers at different stages of pollen development	调控花药组织内活性氧稳态，调控花粉发育相关的多个基因如MS10，AMS等 Regulating redox homeostasis during male gametogenesis；Regulating transcription of genes involved in pollen development，such as MS10，AMS and so on	Salazar-Sarasua et al.,2022
	SlACO2	Solyc12g005860	ACC合成酶 1-aminocyclopropane-1- carboxylicacidoxidase，ACO	未知 Unknown	可能通过乙烯信号调控控制花粉发育 Probably regulating pollen development through ethylene signaling	Secgin et al.,2022
	SlMAPK4	Solyc11g072630	裂原活化蛋白激酶 Mitogen-activated protein kinase 4	仅在成熟花粉粒中表达 Expressed in mature pollens	调控花粉发育相关的碳水化合物代谢基因、信号转导基因。调控花粉壁发育 Involved in pollen wall development. Regulating gens of carbohydrate metabolism and signaling transduction	Wang et al.,2022
	SlBG10	Solyc11g065280	β-1,3-葡聚糖酶β-1,3-glucanase	发育早期在幼蕾期组织内高表达Highly expressed in young stamens at early development stage	具有胼胝质水解酶活性，参与胼胝质代谢 Involved in callus metabolism as callus hydrolase	Pei et al.,2023

基因/位点 Gene/locus	突变位点 Mutation site	分子标记 Molecular marker	参考文献 Reference
PS2	剪接点突变，G ˃ C Mutation in exon-intron splice junction	dCAPS标记，PCR产物用HpyCH4IV酶切 dCAPS marker，the PCR products are digested by restriction enzyme HpyCH4IV Fw：CAAATTGGATGAGAGTTTTGAA Rv：CATTTTACAAGTGTAACAACTTG	Gorguet et al.,2009
MS10	启动子区398 bp 转座子片段插入 Insertion of a 398 bp retrotransposable DNA fragment in promoter	显性标记 Dominant marker Fw：TTCTGTTGTGCGGAATTTGAGATAATAC Rv：CCTACTTCCCTTTCTTCAGAGTTG	Jeong et al.,2014
GPAT6-a	G163R	待开发 To be developed	Petit et al.,2016
MS-15/TM6/TDR6	G110T	InDel标记 InDel marker Fw：TAGAAGAACCCATTGCACTTTGT Rv：CAGAACCTCTCCAACATCATAATC	Cao et al.,2019
MS32	T287*	dCAPS标记，PCR产物用DdeI酶切 dCAPS marker，the PCR products are digested with restriction enzyme DdeI Fw：GATGATTGCGTGCATCTTGG Rv：CGAAGGGTTATCTTGTGATCAAGCGACTT	Liu et al.,2019
MS30	S59I	KASP标记 KASP marker Fw：GAAGGTGACCAAGTTCATGCTTTGACTTACGTAGTAGAAGGGA；GAAGGTCGGAGTCAACGGATTCTTACGTAGTAGAAGGGC Rv：CTTTGCTAGTTCTGGAAAGATGCATG	Wei et al.,2024

Research Progress of Tomato Male Sterility and Its Breeding Application

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靶基因Target gene	基因编号ID	策略Stratedy	材料Genotype	参考文献Reference
LeSUT2	Solyc11g017010	RNAi	Micro-Tom	Hackel et al.,2005
GA20ox1	Solyc03g006880	RNAi	Chico III	Olimpieri et al.,2011
LePro1	Solyc12g044630	RNAi	Moneymaker	Yu & Mandayam,2014
SlMAPK7	Solyc08g081490	RNAi	Micro-Tom	Chen et al.,2016
SlPIN8	Solyc02g087660	RNAi	Micro-Tom	Gan et al.,2019
SlMAPK4	Solyc11g072630	RNAi	Micro-Tom	Wang et al.,2022
Barnase	M14442.1	异源过表达 Heterologous overexpression	Micro-Tom Moneymaker	Cao et al.,2012
SlNCED1	Solyc08g016720	过表达或RNAi Overexpression or RNAi	Jia Bao	Dai et al.,2018
SlMAPK20	Solyc07g056350	RNAi；CRISPR/Cas9基因敲除 RNAi；CRISPR/Cas9 knock out	Micro-Tom	Chen et al.,2018
SlMYB32	Solyc10g055410	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Micro-Tom	Chen et al.,2018
STR	Solyc03g053130	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	TB0993	Du et al.,2020
SlIDA	Solyc05g010000	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Zhongshu 6	Wang et al.,2020
MS10	Solyc02g079810	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Micro-Tom	Liu et al.,2021
PIF3	Solyc01g102300	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Micro-Tom	Yang et al.,2021
SlGLT1	Solyc03g083440	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Micro-Tom	Yang et al.,2021
SlCWIN9	Solyc10g085650	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Micro-Tom	Yang et al.,2021
SlAMS	Solyc08g062780	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Ailsa Craig	Bao et al.,2022
RBOH, RBOHE	Solyc01g099620 Solyc07g042460	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Alisa Craig	Dai et al.,2022
MS1	Solyc04g008420	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Crocker	Gökdemir et al.,2022
SlCMT4	Solyc08g005400.2	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Ailsa Craig	Guo et al.,2022
SlTPD1	Solyc03g097530	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Moneymaker	Salazar-Sarasua et al.,2022
SlBG10	Solyc11g065280	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	Micro-Tom	Pei et al.,2023
MS10	Solyc02g079810	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	TB0993；TB0249	Zhou et al.,2023a
MS15	Solyc02g079810 Solyc02g081340	CRISPR/Cas9基因敲除 CRISPR/Cas9 knock out	TB0993；TB0249	Zhou et al.,2023b

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