Research Progress on the Cytokinin Type-A Response Regulator Gene Family

doi:10.16420/j.issn.0513-353x.2022-0681

Acta Horticulturae Sinica ›› 2023, Vol. 50 ›› Issue (9): 1867-1888.doi: 10.16420/j.issn.0513-353x.2022-0681

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Research Progress on the Cytokinin Type-A Response Regulator Gene Family

LU Jing, WEI Suyun, YIN Tongming, CHEN Yingnan^*()

Key Laboratory of Forest Genetics and Biotechnology of Ministry of Education，College of Forestry，Nanjing Forestry University，Nanjing 210037，China

Received:2023-04-03 Revised:2023-06-13 Online:2023-09-25 Published:2023-09-26
Contact: CHEN Yingnan

Abstract

Abstract:

Type-A response regulators（RRs）are the terminal components of cytokinin signal transduction pathway merely possessing the REC receiving domain.Type-A RRs not only involve in regulating plant growth and development，but also regulating response to abiotic stresses.In this paper，the different aspects of recent studies on gene identification，gene evolutionary relationship，gene expression，and functional characteristics of type-A RRs in herbaceous and woody plants have been reviewed.In addition，the potential regulatory network of type-A RR genes were summarized.This review provided useful information for better understanding the mechanisms by which type-A RRs negatively regulate cytokinin signaling.

Key words: cytokinin, type-A response regulator, expression pattern, gene function, regulatory network

LU Jing, WEI Suyun, YIN Tongming, CHEN Yingnan. Research Progress on the Cytokinin Type-A Response Regulator Gene Family[J]. Acta Horticulturae Sinica, 2023, 50(9): 1867-1888.

Figures/Tables 3

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功能特征 Functional characteristics	物种 Species	基因名 Gene name	亚细胞定位 Subcellular localization	基因表达特性与表型 Gene expression characteristics and phenotype	参考文献 Reference
根生长发育 Root development	拟南芥 Arabidopsis thaliana	ARR5	细胞核 Nucleus	在根顶端分生组织表达量较高 Highly expressed in the apical meristem of roots	To et al.,2004；Dortay et al.,2008
		ARR7	细胞核 Nucleus	过表达株根长；arr7突变体根干细胞发育系统受损 The roots of overexpressing plants were longer；the arr7 mutant has a defective root stem-cell system	Dortay et al.,2008；Müller & Sheen,2008；Lee et al.,2007；Buechel et al.,2010
		ARR15	细胞核 Nucleus	在根维管形成层中表达 Expressed in root vascular cambium Arr15突变体根干细胞发育系统受损 The arr15 mutant has a defective root stem-cell system	Kiba et al.,2002,2003；Dortay et al.,2008；Müller & Sheen,2008；Buechel et al.,2010
		ARR16	细胞质 Cytoplasm	在根内皮层表达 Expressed in root endodermis Arr16/17突变体根尖失去向水性 The root tips of arr16/17 mutant lost hydrotropism	Kiba et al.,2002；Dortay et al.,2008；Chang et al.,2019
		ARR17	-		Kiba et al.,2002；Dortay et al.,2008；Chang et al.,2019
	水稻 Oryza sativa	OsRR2	-	在WOX11突变水稻表达上调；在WOX11过表达植株表达下调；在ren1-D显性突变体水稻表达下调 Upregulated in WOX11 mutant rice plants；downregulated in WOX11 overexpressing plants； downregulated in ren1-D dominant mutant	Zhao et al.,2009； Gao et al.,2014
	水稻 Oryza sativa	OsRR6	-	过表达根系发育不良，不定根数量改变 The overexpressing plants caused defects in the root system and altered adventitious root numbers	Bhaskar et al.,2021
	番茄 Solanum lycopersicum	SIRR4	-	仅在侧根中特异表达 Specifically expressed in lateral roots	Gupta et al.,2013
	大豆 Gylcine max	GmRR06	-	在根中表达量最高 Highest expressed in roots	Le et al.,2011
	毛果杨 Populus trichocarpa	PtRR1	细胞核 Nucleus	在根中表达 Expressed in roots	Ramírez-Carvajal et al.,2008
下胚轴伸长 Hypocotyl elongation	拟南芥 A. thaliana	ARR3	细胞核 Nucleus	arr3/4双突变体在黑暗中生物钟延长，在白光和光/暗周期中下胚轴长 The arr3/4 double mutant has a lengthened circadian in the absence of light，and longer hypocotyls in both white light and light/dark cycles	Salomé et al.,2006；Dortay et al.,2008
	拟南芥 A. thaliana	ARR4	细胞质 Cytoplasm		Salomé et al.,2006；Dortay et al.,2008
	水稻 O. sativa	OsRR6	-	过表达下胚轴生长受到抑制 The hypocotyl growth in overexpressing plants were inhibited	Bhaskar et al.,2021
茎生长发育 Stem development	拟南芥 A. thaliana	ARR4	细胞质 Cytoplasm	在茎中高表达 Highly expressed in stems 过表达芽分化速度快 The overexpressing plants promoted shoot formation	Osakabe et al.,2002；To et al.,2004；Dortay et al.,2008
		ARR5	细胞核 Nucleus	在茎部顶端分生组织中高表达 Highly expressed in the apical meristem of stems	D’ Agostino et al.,2000；Dortay et al.,2008
		ARR7	细胞核 Nucleus	arr7突变体茎尖分生组织发育快 The arr7 mutants have faster development of shoot apical meristem	Zhao et al.,2010
		ARR8	细胞核 Nucleus	过表达芽分化慢 The overexpressing plants inhibited shoot formation	Dortay et al.,2008；Osakabe et al.,2002
		ARR15	细胞核 Nucleus	arr15突变体茎尖分生组织发育快 the arr15 mutants have faster development of apical meristem of stems	Zhao et al.,2010
	水稻 O. sativa	OsRR5	-	在SL相关d53突变体茎中高表达 High expressed in stem of SL-related d53 mutant	Duan et al.,2019
	番茄 S. lycopersicum	SIRR4	-	在番茄茎尖分生组织中高表达 Expressed in the apical meristem of stems in tomato	Shani et al.,2010
叶生长发育 Leaf development	拟南芥 A. thaliana	ARR16	细胞质 Cytoplasm	过表达ARR16叶片黄化严重，气孔数量降低；arr16/17双突变体气孔数量增多 The overexpression of ARR16 caused severe leave senescence and stomas number reduction；the arr16/17 double mutants have more stomas	Dortay et al.,2008；Ren et al.,2009；Vatén et al.,2018
	拟南芥 A. thaliana	ARR17	-		Dortay et al.,2008；Ren et al.,2009；Vatén et al.,2018
	水稻 O. sativa	OsRR6	-	过表达叶片持绿长，花青素积累 The overexpressing plants resulted in sustained green leaves and accumulation of anthocyanin in the leaves	Bhaskar et al.,2021
		OsRR9	-	osrr9/10双突变体延缓叶片衰老 The osrr9/10 double mutant delayed leaf senescence	Wang et al.,2019
		OsRR10	-		Wang et al.,2019
	玉米 Zea mays	ZmRR3	细胞质 Cytoplasm	Mu转座子插入突变改变叶序 Mutator insection caused opposite phyllotaxy	Asakura et al.,2003；Giulini et al.,2004
	番茄 S. lycopersicum	SlRR4	-	在幼叶原基中高表达 Highly expressed in the young leave primordia	Shani et al.,2010
	番茄 S. lycopersicum	SlRR7	-	在幼叶原基中高表达 Highly expressed in the young leave primordia	Shani et al.,2010
芽生长发育 Shoot development	拟南芥 A. thaliana	ARR4	细胞核 Nucleus	过表达芽分化数量多 The overexpressing plants increased the number of shoot formation	Osakabe et al.,2002；Dortay et al.,2008
		ARR8	细胞核 Nucleus	过表达芽分化少 The overexpressing plants decreased the number of shoot formation	Osakabe et al.,2002；Dortay et al.,2008
		ARR15	细胞核 Nucleus	过表达愈伤组织少，芽分化少 The overexpressing plants decreased the formation of callus and shoot	Kiba et al., 2003；Dortay et al.,2008
	大豆 Glycine max	GmRR11	-		Mochida et al.,2010
	苹果 Malus × domestica	MdRR9	-	过表达腋芽和分枝数量显著增加。 The overexpressing plants have more axillary buds and branches	陈皓,2021
	海岸松 Pinus pinaster	PipsRRA1	-	在不定芽中高表达 High expressed in adventitious callogenesis	Alvarez et al.,2012
花发育 Flower development	拟南芥 A. thaliana	ARR7	细胞核 Nucleus	过表达导致花发育异常 The overexpressing plants caused defective flower development	Dortay et al.,2008；Leibfried et al.,2005
	拟南芥 A. thaliana	ARR16	细胞质 Nucleus	在雌核发育的7、8、9和12阶段表达 Expressed during gynoecia development at stage 7，8，9，12	Dortay et al.,2008；Reyes-Olalde et al.,2017
	水稻 O. sativa	OsRR1	-	过表达花期长 The overexpressing plants have a longer flowering time	Cho et al.,2016
		OsRR6	-	过表达圆锥花序发育不良 The overexpressing plants have defective panicle development	Hirose et al.,2007
		OsRR9	-	Osrr9/10双突变体小穗数减少 The number of spikelets in osrr9/10 double mutants was reduced	Wang et al.,2019
		OsRR10	-		Wang et al.,2019
	玉米 Z. mays	ZmRR1 ~ ZmRR2	细胞质 Cytoplasm	在雄花序中表达 Expressed in tassels	Asakura et al.,2003；Sakakibara et al.,1999
	玉米 Z. mays	ZmRR3	细胞质 Cytoplasm	在雌花序中表达 Expressed in ears	Asakura et al.,2003；Sakakibara et al.,1999
	香脂杨 P. balsamifera	PbRR9	-	仅有的性别特异性甲基化基因 Only sex-specific methylation gene	Bräutigam et al.,2017；Cronk et al.,2020
	毛果杨 P. trichocarpa	ARR17	-		Müller et al.,2020
	欧洲山杨 P. tremula	ARR17	-	敲除导致雌株开雄花 Knockout resulted in the production of male flowers in female plants	Müller et al.,2020
	美洲黑杨 P. deltoides	FERR	细胞质 Cytoplasm	仅在雌花原基中高表达 Highly expressed in female flower primordia 过表达促进雌蕊发育 The overexpressing plants promoted carpel development	Xue et al.,2020
	红皮柳 Salix purpurea	GATA15	-	在含有雌花原基的茎尖中特异表达 Specifically expressed in shoot tips containing floral primordia	Hyden et al.,2021
	红皮柳 Salix purpurea	SpRR9	-	在雌株葇夷花序中高表达 Highly expressed in female catkins	Zhou et al.,2020
果实生长发育 Fruit development	番茄 S. lycopersicum	SlRR1	-	在果实发育IG2阶段高表达，在成熟阶段低表达 Highly expressed during fruit developmental stage IG2，and down-regulated during ripening phase	Bianchetti et al.,2017
		SlRR3	-
		SlRR4	-
		SlRR7	-
	苹果 M. domestica	MdRR5	-	在果实脱落前乙烯增加时表达下降 Expression decreased when ethylene increased before fruit shedding	Cin et al.,2009
种子生长发育 Seed development	玉米 Z. mays	ZmTCRR-1	细胞质 Cytoplasm	在未成熟胚乳转移细胞层中特异表达 Specifically expressed in the immature endosperm transfer-cell layer	Asakura et al.,2003；Muñiz et al.,2006
种子生长发育 Seed development	小麦 Triticum aestivum	TaARR9	-	在种子中特异性表达 Specifically expressed in wheat seeds	Tuan et al.,2019
非生物胁迫敏感性 Sensitivity to abiotic stress	拟南芥 A. thaliana	ARR5	细胞核 Nucleus	过表达耐寒 The overexpressing plants increased freezing tolerance.	Dortay et al.,2008；Shi et al.,2012；Huang et al.,2017
		ARR6	细胞核 Nucleus	下调可以提高对氧化应激的耐受性。 Downregulation can enhance the tolerance to oxidative stress	Dortay et al.,2008；Zwack et al.,2016
		ARR7	细胞核 Nucleus	过表达耐寒 The overexpressing plants increased freezing tolerance	Dortay et al.,2008；Jeon et al.,2010
		ARR15	细胞核 Nucleus	过表达耐寒 The overexpressing plants increased freezing tolerance	Dortay et al.,2008；Shi et al.,2012
	水稻 O. sativa	OsRR6	-	过表达在NaCl和甘露醇种子萌发处理下率高 The overexpressing plants have higher seed germination rates with the respective treatment of NaCl and mannitol treatment	Bhaskar et al.,2021
		OsRR9	-	在47 ℃高温下表达上调 Up-regulated at a high temperature of 47 ℃ osrr9/osrr10双突变体在盐胁迫下叶片不易蜷缩 osrr9/osrr10 double mutants reduced leaf curling under salinity stress	Wang et al.,2019
		OsRR10	-	在渗透胁迫和盐胁迫下均表达下调 Down-regulated under osmotic stress and salinity stress	Wang et al.,2019
	玉米 Z.mays	ZmRR1	细胞质 Cytoplasm	在低温处理下，过表达植抗性强，而突变体抗性差 The overexpressing plants exhibited strong resistance while the mutants showed weak resistance under low-temperature treatment	Asakura et al.,2003；Zeng et al.,2021
	白菜 Brassica rapa	BrRR1	-	在盐胁迫处理下，BrRR19基因表达上调；在ABA处理下，BrRR3和BrRR17表达下调；在反式玉米素处理下BrRR1/3/7/8/12/17/19表达上调；在根瘤感染初期BrRR1/6表达上调、在根瘤扩张期，BrRR4表达上调 BrRR19 was upregulated under salt stress；while BrRR3 and BrRR17 were downregulated under ABA treatment；BrRR1/3/7/8/12/17/19 were upregulated under t-zeatin treatment；BrRR1/6 were upregulated during the primary phase of clubroot infection，while BrRR4 was upregulated during the gall enlargement phase	Liu et al.,2014；Laila et al.,2020
		BrRR3	-
		BrRR4	-
		BrRR6	-
		BrRR7	-
		BrRR8	-
		BrRR12	-
		BrRR17	-
		BrRR19	-
	番茄 S. lycopersicum	SIRR5	-	在抗晚疫病的番茄中表达下调，而在晚疫病敏感的番茄中上调Down-regulated in resistant tomato，while up-regulated in sensitive tomato	Cui et al.,2017
	百脉根 Lotus japonicus	LjaRRa1 ~ LjaRRa6	-	在反式玉米素处理后表达上调 Up-regulated after t-zeatin treatment	Ishida et al.,2009
	苹果 M. domestica	MdRR3	-	在6-BA处理后表达上调 Up-regulated after 6-BA treatment	Li et al.,2017
	苹果 M. domestica	MdRR14	-	在6-BA处理后表达上调 Up-regulated after 6-BA treatment	Li et al.,2017
	银灰杨 P. × canescens	PtaRR3	-	干旱胁迫下在杨树的茎的初生木质部中表达下调 Down-regulated in the primary xylem of poplar stems under drought stress	Paul et al.,2017

功能特征 Functional characteristics	物种 Species	基因名 Gene name	亚细胞定位 Subcellular localization	基因表达特性与表型 Gene expression characteristics and phenotype	参考文献 Reference
根生长发育 Root development	拟南芥 Arabidopsis thaliana	ARR5	细胞核 Nucleus	在根顶端分生组织表达量较高 Highly expressed in the apical meristem of roots	To et al.,2004；Dortay et al.,2008
		ARR7	细胞核 Nucleus	过表达株根长；arr7突变体根干细胞发育系统受损 The roots of overexpressing plants were longer；the arr7 mutant has a defective root stem-cell system	Dortay et al.,2008；Müller & Sheen,2008；Lee et al.,2007；Buechel et al.,2010
		ARR15	细胞核 Nucleus	在根维管形成层中表达 Expressed in root vascular cambium Arr15突变体根干细胞发育系统受损 The arr15 mutant has a defective root stem-cell system	Kiba et al.,2002,2003；Dortay et al.,2008；Müller & Sheen,2008；Buechel et al.,2010
		ARR16	细胞质 Cytoplasm	在根内皮层表达 Expressed in root endodermis Arr16/17突变体根尖失去向水性 The root tips of arr16/17 mutant lost hydrotropism	Kiba et al.,2002；Dortay et al.,2008；Chang et al.,2019
		ARR17	-		Kiba et al.,2002；Dortay et al.,2008；Chang et al.,2019
	水稻 Oryza sativa	OsRR2	-	在WOX11突变水稻表达上调；在WOX11过表达植株表达下调；在ren1-D显性突变体水稻表达下调 Upregulated in WOX11 mutant rice plants；downregulated in WOX11 overexpressing plants； downregulated in ren1-D dominant mutant	Zhao et al.,2009； Gao et al.,2014
	水稻 Oryza sativa	OsRR6	-	过表达根系发育不良，不定根数量改变 The overexpressing plants caused defects in the root system and altered adventitious root numbers	Bhaskar et al.,2021
	番茄 Solanum lycopersicum	SIRR4	-	仅在侧根中特异表达 Specifically expressed in lateral roots	Gupta et al.,2013
	大豆 Gylcine max	GmRR06	-	在根中表达量最高 Highest expressed in roots	Le et al.,2011
	毛果杨 Populus trichocarpa	PtRR1	细胞核 Nucleus	在根中表达 Expressed in roots	Ramírez-Carvajal et al.,2008
下胚轴伸长 Hypocotyl elongation	拟南芥 A. thaliana	ARR3	细胞核 Nucleus	arr3/4双突变体在黑暗中生物钟延长，在白光和光/暗周期中下胚轴长 The arr3/4 double mutant has a lengthened circadian in the absence of light，and longer hypocotyls in both white light and light/dark cycles	Salomé et al.,2006；Dortay et al.,2008
	拟南芥 A. thaliana	ARR4	细胞质 Cytoplasm		Salomé et al.,2006；Dortay et al.,2008
	水稻 O. sativa	OsRR6	-	过表达下胚轴生长受到抑制 The hypocotyl growth in overexpressing plants were inhibited	Bhaskar et al.,2021
茎生长发育 Stem development	拟南芥 A. thaliana	ARR4	细胞质 Cytoplasm	在茎中高表达 Highly expressed in stems 过表达芽分化速度快 The overexpressing plants promoted shoot formation	Osakabe et al.,2002；To et al.,2004；Dortay et al.,2008
		ARR5	细胞核 Nucleus	在茎部顶端分生组织中高表达 Highly expressed in the apical meristem of stems	D’ Agostino et al.,2000；Dortay et al.,2008
		ARR7	细胞核 Nucleus	arr7突变体茎尖分生组织发育快 The arr7 mutants have faster development of shoot apical meristem	Zhao et al.,2010
		ARR8	细胞核 Nucleus	过表达芽分化慢 The overexpressing plants inhibited shoot formation	Dortay et al.,2008；Osakabe et al.,2002
		ARR15	细胞核 Nucleus	arr15突变体茎尖分生组织发育快 the arr15 mutants have faster development of apical meristem of stems	Zhao et al.,2010
	水稻 O. sativa	OsRR5	-	在SL相关d53突变体茎中高表达 High expressed in stem of SL-related d53 mutant	Duan et al.,2019
	番茄 S. lycopersicum	SIRR4	-	在番茄茎尖分生组织中高表达 Expressed in the apical meristem of stems in tomato	Shani et al.,2010
叶生长发育 Leaf development	拟南芥 A. thaliana	ARR16	细胞质 Cytoplasm	过表达ARR16叶片黄化严重，气孔数量降低；arr16/17双突变体气孔数量增多 The overexpression of ARR16 caused severe leave senescence and stomas number reduction；the arr16/17 double mutants have more stomas	Dortay et al.,2008；Ren et al.,2009；Vatén et al.,2018
	拟南芥 A. thaliana	ARR17	-		Dortay et al.,2008；Ren et al.,2009；Vatén et al.,2018
	水稻 O. sativa	OsRR6	-	过表达叶片持绿长，花青素积累 The overexpressing plants resulted in sustained green leaves and accumulation of anthocyanin in the leaves	Bhaskar et al.,2021
		OsRR9	-	osrr9/10双突变体延缓叶片衰老 The osrr9/10 double mutant delayed leaf senescence	Wang et al.,2019
		OsRR10	-		Wang et al.,2019
	玉米 Zea mays	ZmRR3	细胞质 Cytoplasm	Mu转座子插入突变改变叶序 Mutator insection caused opposite phyllotaxy	Asakura et al.,2003；Giulini et al.,2004
	番茄 S. lycopersicum	SlRR4	-	在幼叶原基中高表达 Highly expressed in the young leave primordia	Shani et al.,2010
	番茄 S. lycopersicum	SlRR7	-	在幼叶原基中高表达 Highly expressed in the young leave primordia	Shani et al.,2010
芽生长发育 Shoot development	拟南芥 A. thaliana	ARR4	细胞核 Nucleus	过表达芽分化数量多 The overexpressing plants increased the number of shoot formation	Osakabe et al.,2002；Dortay et al.,2008
		ARR8	细胞核 Nucleus	过表达芽分化少 The overexpressing plants decreased the number of shoot formation	Osakabe et al.,2002；Dortay et al.,2008
		ARR15	细胞核 Nucleus	过表达愈伤组织少，芽分化少 The overexpressing plants decreased the formation of callus and shoot	Kiba et al., 2003；Dortay et al.,2008
	大豆 Glycine max	GmRR11	-		Mochida et al.,2010
	苹果 Malus × domestica	MdRR9	-	过表达腋芽和分枝数量显著增加。 The overexpressing plants have more axillary buds and branches	陈皓,2021
	海岸松 Pinus pinaster	PipsRRA1	-	在不定芽中高表达 High expressed in adventitious callogenesis	Alvarez et al.,2012
花发育 Flower development	拟南芥 A. thaliana	ARR7	细胞核 Nucleus	过表达导致花发育异常 The overexpressing plants caused defective flower development	Dortay et al.,2008；Leibfried et al.,2005
	拟南芥 A. thaliana	ARR16	细胞质 Nucleus	在雌核发育的7、8、9和12阶段表达 Expressed during gynoecia development at stage 7，8，9，12	Dortay et al.,2008；Reyes-Olalde et al.,2017
	水稻 O. sativa	OsRR1	-	过表达花期长 The overexpressing plants have a longer flowering time	Cho et al.,2016
		OsRR6	-	过表达圆锥花序发育不良 The overexpressing plants have defective panicle development	Hirose et al.,2007
		OsRR9	-	Osrr9/10双突变体小穗数减少 The number of spikelets in osrr9/10 double mutants was reduced	Wang et al.,2019
		OsRR10	-		Wang et al.,2019
	玉米 Z. mays	ZmRR1 ~ ZmRR2	细胞质 Cytoplasm	在雄花序中表达 Expressed in tassels	Asakura et al.,2003；Sakakibara et al.,1999
	玉米 Z. mays	ZmRR3	细胞质 Cytoplasm	在雌花序中表达 Expressed in ears	Asakura et al.,2003；Sakakibara et al.,1999
	香脂杨 P. balsamifera	PbRR9	-	仅有的性别特异性甲基化基因 Only sex-specific methylation gene	Bräutigam et al.,2017；Cronk et al.,2020
	毛果杨 P. trichocarpa	ARR17	-		Müller et al.,2020
	欧洲山杨 P. tremula	ARR17	-	敲除导致雌株开雄花 Knockout resulted in the production of male flowers in female plants	Müller et al.,2020
	美洲黑杨 P. deltoides	FERR	细胞质 Cytoplasm	仅在雌花原基中高表达 Highly expressed in female flower primordia 过表达促进雌蕊发育 The overexpressing plants promoted carpel development	Xue et al.,2020
	红皮柳 Salix purpurea	GATA15	-	在含有雌花原基的茎尖中特异表达 Specifically expressed in shoot tips containing floral primordia	Hyden et al.,2021
	红皮柳 Salix purpurea	SpRR9	-	在雌株葇夷花序中高表达 Highly expressed in female catkins	Zhou et al.,2020
果实生长发育 Fruit development	番茄 S. lycopersicum	SlRR1	-	在果实发育IG2阶段高表达，在成熟阶段低表达 Highly expressed during fruit developmental stage IG2，and down-regulated during ripening phase	Bianchetti et al.,2017
		SlRR3	-
		SlRR4	-
		SlRR7	-
	苹果 M. domestica	MdRR5	-	在果实脱落前乙烯增加时表达下降 Expression decreased when ethylene increased before fruit shedding	Cin et al.,2009
种子生长发育 Seed development	玉米 Z. mays	ZmTCRR-1	细胞质 Cytoplasm	在未成熟胚乳转移细胞层中特异表达 Specifically expressed in the immature endosperm transfer-cell layer	Asakura et al.,2003；Muñiz et al.,2006
种子生长发育 Seed development	小麦 Triticum aestivum	TaARR9	-	在种子中特异性表达 Specifically expressed in wheat seeds	Tuan et al.,2019
非生物胁迫敏感性 Sensitivity to abiotic stress	拟南芥 A. thaliana	ARR5	细胞核 Nucleus	过表达耐寒 The overexpressing plants increased freezing tolerance.	Dortay et al.,2008；Shi et al.,2012；Huang et al.,2017
		ARR6	细胞核 Nucleus	下调可以提高对氧化应激的耐受性。 Downregulation can enhance the tolerance to oxidative stress	Dortay et al.,2008；Zwack et al.,2016
		ARR7	细胞核 Nucleus	过表达耐寒 The overexpressing plants increased freezing tolerance	Dortay et al.,2008；Jeon et al.,2010
		ARR15	细胞核 Nucleus	过表达耐寒 The overexpressing plants increased freezing tolerance	Dortay et al.,2008；Shi et al.,2012
	水稻 O. sativa	OsRR6	-	过表达在NaCl和甘露醇种子萌发处理下率高 The overexpressing plants have higher seed germination rates with the respective treatment of NaCl and mannitol treatment	Bhaskar et al.,2021
		OsRR9	-	在47 ℃高温下表达上调 Up-regulated at a high temperature of 47 ℃ osrr9/osrr10双突变体在盐胁迫下叶片不易蜷缩 osrr9/osrr10 double mutants reduced leaf curling under salinity stress	Wang et al.,2019
		OsRR10	-	在渗透胁迫和盐胁迫下均表达下调 Down-regulated under osmotic stress and salinity stress	Wang et al.,2019
	玉米 Z.mays	ZmRR1	细胞质 Cytoplasm	在低温处理下，过表达植抗性强，而突变体抗性差 The overexpressing plants exhibited strong resistance while the mutants showed weak resistance under low-temperature treatment	Asakura et al.,2003；Zeng et al.,2021
	白菜 Brassica rapa	BrRR1	-	在盐胁迫处理下，BrRR19基因表达上调；在ABA处理下，BrRR3和BrRR17表达下调；在反式玉米素处理下BrRR1/3/7/8/12/17/19表达上调；在根瘤感染初期BrRR1/6表达上调、在根瘤扩张期，BrRR4表达上调 BrRR19 was upregulated under salt stress；while BrRR3 and BrRR17 were downregulated under ABA treatment；BrRR1/3/7/8/12/17/19 were upregulated under t-zeatin treatment；BrRR1/6 were upregulated during the primary phase of clubroot infection，while BrRR4 was upregulated during the gall enlargement phase	Liu et al.,2014；Laila et al.,2020
		BrRR3	-
		BrRR4	-
		BrRR6	-
		BrRR7	-
		BrRR8	-
		BrRR12	-
		BrRR17	-
		BrRR19	-
	番茄 S. lycopersicum	SIRR5	-	在抗晚疫病的番茄中表达下调，而在晚疫病敏感的番茄中上调Down-regulated in resistant tomato，while up-regulated in sensitive tomato	Cui et al.,2017
	百脉根 Lotus japonicus	LjaRRa1 ~ LjaRRa6	-	在反式玉米素处理后表达上调 Up-regulated after t-zeatin treatment	Ishida et al.,2009
	苹果 M. domestica	MdRR3	-	在6-BA处理后表达上调 Up-regulated after 6-BA treatment	Li et al.,2017
	苹果 M. domestica	MdRR14	-	在6-BA处理后表达上调 Up-regulated after 6-BA treatment	Li et al.,2017
	银灰杨 P. × canescens	PtaRR3	-	干旱胁迫下在杨树的茎的初生木质部中表达下调 Down-regulated in the primary xylem of poplar stems under drought stress	Paul et al.,2017

Research Progress on the Cytokinin Type-A Response Regulator Gene Family

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