油菜素内酯调控植物株高的作用机制及应用

doi:10.16420/j.issn.0513-353x.2025-0519

摘要/Abstract

摘要：

株高是决定植物株型结构、抗倒伏能力及产量的关键性状，油菜素内酯（brassinosteroid，BR）对株高调控起重要作用。综述了BR调控植物株高的作用：在生理层面上，BR通过调控细胞伸长与细胞分裂过程影响株高，作用机制涉及水分吸收、细胞壁松弛及微管稳定性等方面；在分子机制上，BR通过生物合成基因（如DWF4、CPD、DET2、ROT3、CYP90D1和CYP85A1/2）及信号转导基因（如BRI1、BAK1、BSU1、BIN2、BZR1和BES1）精确调控株高，并与赤霉素、生长素、茉莉酸、脱落酸、乙烯、独脚金内酯等多种激素相互作用，形成复杂的株高调控网络。本文中还总结了BR在矮化育种中的应用，BR相关矮化种质在植物株型改良与产量提升方面显示出巨大潜力。最后，分析当前研究中存在的问题并进行展望，以期为植物株高调控机制的深入研究和矮化育种实践提供参考。

关键词: 油菜素内酯, 株高, 调控机制, 矮化育种

Abstract:

Plant height is a key trait that determines plant architecture，lodging resistance，and yield. Brassinosteroid（BR）plays a crucial role in regulating plant height. This review summarizes recent advances in understanding how BR regulates plant height. At the physiological level，BR modulates cell elongation and division by affecting water uptake，cell wall modification，and microtubule stability. At the molecular level，BR precisely controls plant height through key biosynthesis genes such as DWF4，CPD，DET2，ROT3，CYP90D1，and CYP85A1/2，as well as signal transduction genes including BRI1，BAK1，BSU1，BIN2，BZR1，and BES1. During their functioning，BR crosstalks with gibberellin，auxin，jasmonate，abscisic acid，ethylene，and strigolactone，forming a complex regulatory network for plant height. The application of BR in dwarf breeding is also discussed，BR-related dwarf germplasm demonstrates significant potential in improving plant architecture and increasing yield. Finally，the problems existing in the current research are analyzed and prospects are put forward. The key research directions highlighted in this review will provide a a reference for the in-depth study on the regulatory mechanism of plant height and the practice of dwarf breeding.

Key words: brassinosteroid, plant height, regulatory mechanisms, dwarf breeding

吕鑫阳, 郭悦, 冯明涵, 祝朋芳, 冯馨. 油菜素内酯调控植物株高的作用机制及应用[J]. 园艺学报, 2026, 53(5): 1535-1552.

LÜ Xinyang, GUO Yue, FENG Minghan, ZHU Pengfang, FENG Xin. Research Progress on the Mechanism and Application of Brassinosteroid in Regulating Plant Height[J]. Acta Horticulturae Sinica, 2026, 53(5): 1535-1552.

图/表 3

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通路 Pathway	拟南芥 Arabidopsis thaliana		水稻 Oryza sativa		玉米 Zea mays		番茄 Solanum lycopersicum
通路 Pathway	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID
BR合成 BR biosynthesis	AtDWF4	AT3G50660	OsDWARF4	Os03g0227700	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtDWF4	AT3G50660	OsD11	Os04g0469800	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtCPD	AT5G05690	OsCPD1	Os11g0143200			SlCPD	Solyc06g051750
	AtCPD	AT5G05690	OsCPD2	Os12g0139300			SlCPD	Solyc06g051750
	AtDET2	AT2G38050	OsDET2	Os01g0851600	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtDET2	AT2G38050	OsDET2	Os11g0184100	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtROT3	AT4G36380					SlROT3	Solyc02g084740
	AtCYP90D1	AT3G13730					SlCYP90D1	Solyc03g121510
	AtCYP85A1	AT5G38970	OsCYP85A1	Os03g0602300	ZmBRD1	GRMZM2G103773	SlDWARF	Solyc02g089160
	AtCYP85A2	AT3G30180
BR信号转导 BR signaling	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1a	GRMZM2G048294	SlBRI1	Solyc04g051510
	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1b	GRMZM2G449830	SlBRI1	Solyc04g051510
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc10g047140
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc01g104970
	AtBSU1	AT1G03445	OsPPKL1	Os03g0646900			SlBSU1	Solyc09g074320
	AtBSL1	AT4G03080	OsPPKL2	Os05g0144400
	AtBSL2	AT1G08420	OsPPKL1	Os03g0646900
	AtBSL3	AT2G27210	OsPPKL3	Os12g0617900
BR信号转导 BR signaling	AtBIN2	AT4G18710	OsGSK1	Os01g0205700	ZmGSK2	GRMZM2G472625	SlBIN2.1	Solyc02g072300
			OsGSK2	Os05g0207500			SlBIN2.2	Solyc07g055200
			OsGSK2	Os05g0207500			SlBIN2.3	Solyc03g006070
	AtBZR1	AT1G75080	OsBZR1	Os07g0580500	ZmBZR1	GRMZM5G852801	SlBZR1	Solyc12g089040
	AtBES1	AT1G19350	OsBZR2	Os01g0203000	ZmBES1	GRMZM2G102514	SlBES1	Solyc04g079980

通路 Pathway	拟南芥 Arabidopsis thaliana		水稻 Oryza sativa		玉米 Zea mays		番茄 Solanum lycopersicum
通路 Pathway	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID	基因名 Gene name	基因ID Gene ID
BR合成 BR biosynthesis	AtDWF4	AT3G50660	OsDWARF4	Os03g0227700	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtDWF4	AT3G50660	OsD11	Os04g0469800	ZmDWF4	GRMZM2G065635	SlCYP90B3	Solyc02g085360
	AtCPD	AT5G05690	OsCPD1	Os11g0143200			SlCPD	Solyc06g051750
	AtCPD	AT5G05690	OsCPD2	Os12g0139300			SlCPD	Solyc06g051750
	AtDET2	AT2G38050	OsDET2	Os01g0851600	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtDET2	AT2G38050	OsDET2	Os11g0184100	ZmDET2	GRMZM2G449033	SlDET2	Solyc10g086500
	AtROT3	AT4G36380					SlROT3	Solyc02g084740
	AtCYP90D1	AT3G13730					SlCYP90D1	Solyc03g121510
	AtCYP85A1	AT5G38970	OsCYP85A1	Os03g0602300	ZmBRD1	GRMZM2G103773	SlDWARF	Solyc02g089160
	AtCYP85A2	AT3G30180
BR信号转导 BR signaling	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1a	GRMZM2G048294	SlBRI1	Solyc04g051510
	AtBRI1	AT4G39400	OsBRI1	Os01g0718300	ZmBRI1b	GRMZM2G449830	SlBRI1	Solyc04g051510
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc10g047140
	AtBAK1	AT4G33430	OsBAK1	Os08g0174700	ZmBAK1	GRMZM2G145720	SlBAK1	Solyc01g104970
	AtBSU1	AT1G03445	OsPPKL1	Os03g0646900			SlBSU1	Solyc09g074320
	AtBSL1	AT4G03080	OsPPKL2	Os05g0144400
	AtBSL2	AT1G08420	OsPPKL1	Os03g0646900
	AtBSL3	AT2G27210	OsPPKL3	Os12g0617900
BR信号转导 BR signaling	AtBIN2	AT4G18710	OsGSK1	Os01g0205700	ZmGSK2	GRMZM2G472625	SlBIN2.1	Solyc02g072300
			OsGSK2	Os05g0207500			SlBIN2.2	Solyc07g055200
			OsGSK2	Os05g0207500			SlBIN2.3	Solyc03g006070
	AtBZR1	AT1G75080	OsBZR1	Os07g0580500	ZmBZR1	GRMZM5G852801	SlBZR1	Solyc12g089040
	AtBES1	AT1G19350	OsBZR2	Os01g0203000	ZmBES1	GRMZM2G102514	SlBES1	Solyc04g079980

通路 Pathway	基因名称 Gene name	功能丧失型突变体 Loss-of- function mutant	功能丧失型突变体表型 Phenotype of the loss-of- function mutant	功能获得型突变体/过表达植株 Gain-of-function mutant/ Overexpression plant	功能获得型突变体/过表达植株表型 Phenotype of the gain-of-function mutant or overexpression plant	参考文献 Reference
BR合成 BR biosynthesis	AtDWF4	dwf4	矮化，深绿色，叶片小而圆，叶柄短，莲座叶数量增加，顶端优势减弱，雄性不育，开花延迟 Dwarf，dark green，small and rounded leaves，short petioles，increased number of rosette leaves，reduced apical dominance，male sterility，delayed flowering	AOD4	株高增加，分枝和角果数量增多，叶片和叶柄伸长 Increased plant height，increased branching and silique number，elongated leaves and petioles	Azpiroz et al.,1998； Choe et al.,2001
	AtCPD	cpd	矮化，叶片小而圆，花序茎和花器官缩短，雄性不育 Dwarf，small and rounded leaves，shortened inflorescence stems，and flower organs，male sterility	cpd comp.	将CPD cDNA在cpd 突变体中表达，突变体恢复了野生型的表型，甚至一部分比野生型叶片更大、分枝更多 Overexpression of CPD cDNA in the cpd mutant restored the wild-type phenotype；some lines exhibited even larger leaves and more branching than wild-type	Szekeres et al.,1996； Zhiponova et al.,2013
	AtDET2	det2	矮化，深绿色，叶片小，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，small leaves，reduced apical dominance，reduced male fertility，delayed flowering			Chory et al.,1991； Li et al.,1996
	AtROT3	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation	A-1（ROT3 OE）	叶片和花器官伸长 Elongated leaves and flower organs	Kim et al.,1999； Kim et al.,2005a
	AtCYP90D1	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation			Kim et al.,1999； Kim et al.,2005a
	AtCYP85A1	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A1	表型无变化 Do not cause any phenotypic changes	Kim et al.,2005b； Nomura et al.,2005
	AtCYP85A2	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A2	莲座叶更大，叶柄更长，茎更高且多分枝，角果更大，花梗更长 Larger rosette leaves，longer petioles，taller stems with increased branching，larger siliques，longer pedicels	Kim et al.,2005b； Nomura et al.,2005
BR信号转导 BR signaling	AtBRI1	bri1	严重矮化，深绿色，叶片厚，顶端优势减弱，雄性不育 Severe dwarfism，dark green，thickened leaves，reduced apical dominance，male sterility	BRI1-GFP	叶柄伸长 Elongated petioles	Clouse,1996； Wang et al.,2001； Blanco- Touriñán et al.,2024
	AtBAK1	bak1-1	半矮化，莲座叶小而圆，叶柄短 Semi-dwarf，small and rounded rosette leaves，short petioles	bak1-1D	茎、叶和叶柄伸长 Elongated stems，leaves，and petioles	Li et al.,2002
	AtBSU1	bsu1;bsl1/BSL2,3-amiRNA	严重矮化 Severe dwarfism	BSU1-YFP/bri1-116	在bri1-116中过表达BSU1，部分挽救了其矮化表型 Overexpression of BSU1 in bri1-116 partially rescued the dwarf phenotype	Kim et al.,2009； Kim et al.,2018
	AtBSL1			BSL1-YFP/bri1-5	部分挽救了其矮化表型 Overexpression of BSL1 in bri1-5 partially rescued the dwarf phenotype
	AtBSL2			BSL2-YFP	叶柄伸长 Elongated petioles
	AtBSL3
	AtBIN2	Triple GSK3 mutant	下胚轴显著伸长 Significantly elongated hypocoty	bin2-1	矮化，深绿色，叶片卷曲，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，curled leaves，reduced apical dominance，reduced male fertility，delayed flowering	Li et al.,2001； Vert & Chory,2006
	AtBZR1	BES1 RNAi	半矮化 Semi-dwarf	bzr1-1D	在黑暗中，下胚轴伸长；在光下，半矮化，呈深绿色，叶片宽，叶柄短，叶片数量增加，开花延迟 In darkness：elongated hypocotyl；In light：semi- dwarf，dark green，wide leaves，short petioles，increased leaf number，delayed flowering	Wang et al.,2002； Yin et al.,2005
	AtBES1	BES1 RNAi	半矮化 Semi-dwarf	bes1-D	完全挽救bri1的矮化表型，叶柄长而弯曲，叶片卷曲，过度的茎伸长 Fully rescued the dwarf phenotype of bri1，long and curved petioles，curled leaves，excessive stem elongation	Yin et al.,2002； Yin et al.,2005

通路 Pathway	基因名称 Gene name	功能丧失型突变体 Loss-of- function mutant	功能丧失型突变体表型 Phenotype of the loss-of- function mutant	功能获得型突变体/过表达植株 Gain-of-function mutant/ Overexpression plant	功能获得型突变体/过表达植株表型 Phenotype of the gain-of-function mutant or overexpression plant	参考文献 Reference
BR合成 BR biosynthesis	AtDWF4	dwf4	矮化，深绿色，叶片小而圆，叶柄短，莲座叶数量增加，顶端优势减弱，雄性不育，开花延迟 Dwarf，dark green，small and rounded leaves，short petioles，increased number of rosette leaves，reduced apical dominance，male sterility，delayed flowering	AOD4	株高增加，分枝和角果数量增多，叶片和叶柄伸长 Increased plant height，increased branching and silique number，elongated leaves and petioles	Azpiroz et al.,1998； Choe et al.,2001
	AtCPD	cpd	矮化，叶片小而圆，花序茎和花器官缩短，雄性不育 Dwarf，small and rounded leaves，shortened inflorescence stems，and flower organs，male sterility	cpd comp.	将CPD cDNA在cpd 突变体中表达，突变体恢复了野生型的表型，甚至一部分比野生型叶片更大、分枝更多 Overexpression of CPD cDNA in the cpd mutant restored the wild-type phenotype；some lines exhibited even larger leaves and more branching than wild-type	Szekeres et al.,1996； Zhiponova et al.,2013
	AtDET2	det2	矮化，深绿色，叶片小，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，small leaves，reduced apical dominance，reduced male fertility，delayed flowering			Chory et al.,1991； Li et al.,1996
	AtROT3	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation	A-1（ROT3 OE）	叶片和花器官伸长 Elongated leaves and flower organs	Kim et al.,1999； Kim et al.,2005a
	AtCYP90D1	rot3-4 cyp90d1	严重矮化，下胚轴伸长缺陷 Severe dwarfism，defective hypocotyl elongation			Kim et al.,1999； Kim et al.,2005a
	AtCYP85A1	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A1	表型无变化 Do not cause any phenotypic changes	Kim et al.,2005b； Nomura et al.,2005
	AtCYP85A2	cyp85a1-1/cyp85a2-2	严重矮化 Severe dwarfism	35S-CYP85A2	莲座叶更大，叶柄更长，茎更高且多分枝，角果更大，花梗更长 Larger rosette leaves，longer petioles，taller stems with increased branching，larger siliques，longer pedicels	Kim et al.,2005b； Nomura et al.,2005
BR信号转导 BR signaling	AtBRI1	bri1	严重矮化，深绿色，叶片厚，顶端优势减弱，雄性不育 Severe dwarfism，dark green，thickened leaves，reduced apical dominance，male sterility	BRI1-GFP	叶柄伸长 Elongated petioles	Clouse,1996； Wang et al.,2001； Blanco- Touriñán et al.,2024
	AtBAK1	bak1-1	半矮化，莲座叶小而圆，叶柄短 Semi-dwarf，small and rounded rosette leaves，short petioles	bak1-1D	茎、叶和叶柄伸长 Elongated stems，leaves，and petioles	Li et al.,2002
	AtBSU1	bsu1;bsl1/BSL2,3-amiRNA	严重矮化 Severe dwarfism	BSU1-YFP/bri1-116	在bri1-116中过表达BSU1，部分挽救了其矮化表型 Overexpression of BSU1 in bri1-116 partially rescued the dwarf phenotype	Kim et al.,2009； Kim et al.,2018
	AtBSL1			BSL1-YFP/bri1-5	部分挽救了其矮化表型 Overexpression of BSL1 in bri1-5 partially rescued the dwarf phenotype
	AtBSL2			BSL2-YFP	叶柄伸长 Elongated petioles
	AtBSL3
	AtBIN2	Triple GSK3 mutant	下胚轴显著伸长 Significantly elongated hypocoty	bin2-1	矮化，深绿色，叶片卷曲，顶端优势减弱，雄性育性降低，开花延迟 Dwarf，dark green，curled leaves，reduced apical dominance，reduced male fertility，delayed flowering	Li et al.,2001； Vert & Chory,2006
	AtBZR1	BES1 RNAi	半矮化 Semi-dwarf	bzr1-1D	在黑暗中，下胚轴伸长；在光下，半矮化，呈深绿色，叶片宽，叶柄短，叶片数量增加，开花延迟 In darkness：elongated hypocotyl；In light：semi- dwarf，dark green，wide leaves，short petioles，increased leaf number，delayed flowering	Wang et al.,2002； Yin et al.,2005
	AtBES1	BES1 RNAi	半矮化 Semi-dwarf	bes1-D	完全挽救bri1的矮化表型，叶柄长而弯曲，叶片卷曲，过度的茎伸长 Fully rescued the dwarf phenotype of bri1，long and curved petioles，curled leaves，excessive stem elongation	Yin et al.,2002； Yin et al.,2005