植物表皮蜡质生物合成及其分子调控研究进展

doi:10.13304/j.nykjdb.2024.0290

摘要/Abstract

摘要：

表皮蜡质是覆盖在陆地植物地上部分最外层的一层疏水性保护屏障，在植物抵御生物和非生物胁迫等方面起重要作用。模式植物表皮蜡质生物合成转运途径已被逐渐揭示，一些结构基因和调控基因相继被发掘和鉴定，但其合成转运的分子调控机制仍不很清楚，且在非模式植物中的研究较少。总结了植物表皮蜡质结构、组分和功能，并阐述了植物表皮蜡质的生物合成和转运机制、相关结构基因的功能，重点梳理了植物表皮蜡质生物合成的转录、转录后、翻译后和表观遗传调控机制，并对今后研究重点进行展望，以期为植物表皮蜡质的后续研究与农业生产实践提供启示。

关键词: 植物, 表皮蜡质, 抗胁迫, 生物合成, 调控机制

Abstract:

The cuticular wax， a hydrophobic protective barrier covering most aerial plant organs， plays crucial roles in defending against biotic and abiotic stresses. Although the cuticular wax biosynthesis and transport pathway has been progressively elucidated in model plants， with the identification of several structural and regulatory genes， the underlying molecular mechanisms remain largely unknown， particularly in non-model plants. The structure， components， and functions of the plant cuticular wax were introduced，and the mechanisms of cuticular wax biosynthesis and transport were outlined， highlighting the roles of associated structural genes. The regulatory mechanisms underlying cuticular wax biosynthesis at the transcriptional， post-transcriptional， post-translational， and epigenetic levels were critically examined. Finally， the major research trends in this field were discussed，which aimed to provide valuable insights for further research on plant cuticular wax and its implications for agricultural production.

Key words: plant, cuticular wax, stress resistance, biosynthesis, regulatory mechanism

中图分类号:

Q946.4

赵雄, 马关柳, 郭通琴, 何国菊, 赵许朋, 刘声传. 植物表皮蜡质生物合成及其分子调控研究进展[J]. 中国农业科技导报, 2025, 27(10): 10-23.

Xiong ZHAO, Guanliu MA, Tongqin GUO, Guoju HE, Xupeng ZHAO, Shengchuan LIU. Recent Advances on Cuticular Wax Biosynthesis and Its Molecular Regulation in Plants[J]. Journal of Agricultural Science and Technology, 2025, 27(10): 10-23.

图/表 6

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物种 Species	基因名 Gene name	功能 Function	参考文献 Reference
拟南芥Arabidopsis thaliana	CER17	超长链酰基辅酶A的n-6去饱和 n-6 desaturation of very long chain acyl-CoAs	［4］
拟南芥Arabidopsis thaliana	LACS1， LACS2， LACS4	蜡质合成过程中重叠 Overlap in wax synthesis	［11］
拟南芥Arabidopsis thaliana	Cytb5R	CER1的辅助因子Cofactor of CER1	［19］
拟南芥Arabidopsis thaliana	WAX2/CER3	醛类合成Synthesis of aldehydes	［20］
拟南芥Arabidopsis thaliana	ABCG11/WBC11，ABCG12/CER5	蜡质转运Cuticular wax transport	［23，27］
拟南芥Arabidopsis thaliana	ACC1	丙二酰辅酶 A 底物合成 Synthesis of malonyl-CoA substrate	［29］
拟南芥Arabidopsis thaliana	KCS3	负调控蜡质合成 Negative regulation of wax biosynthesis	［30］
番茄Solanum lycopersicum	CER1-1	烷烃类合成Biosynthesis of alkanes	［34］
拟南芥Arabidopsis thaliana	KCS1	延伸C26～C30脂肪酸 Extension of C26 ~ C30 fatty acids	［43］
拟南芥Arabidopsis thaliana	CUT1， CER6， KCS5， KCS6	调控超长链脂肪酸合成 Regulation of VLCFA biosynthesis	［44］
拟南芥Arabidopsis thaliana	FATA/FATB	为蜡质合成提供饱和脂肪酸 Provide saturated fatty acids for wax biosynthesis	［45］
拟南芥Arabidopsis thaliana	CER10	超长链脂肪酸合成VLCFA biosynthesis	［46］
拟南芥Arabidopsis thaliana	PAS2/HCD	超长链脂肪酸合成VLCFA biosynthesis	［47］
拟南芥Arabidopsis thaliana	KCS20， KCS2	超长链脂肪酸扩展至C22 Extende VLCFAs to C22	［48］
拟南芥Arabidopsis thaliana	LACS1/CER8，LCAS2， LCAS4	超长链脂肪酸C20～C30 合成 Synthetase activity for VLCFAs C20-C30	［49］
拟南芥Arabidopsis thaliana	CER2， CER2-Like1/2	大于C28脂肪酸的延伸 Extension of fatty acids beyond C28	［50］
拟南芥Arabidopsis thaliana	KCS9	延伸C22～C24脂肪酸 Extension of C22～C24 fatty acids	［51］
拟南芥Arabidopsis thaliana	CER1， CER22	烷烃类合成Biosynthesis of alkanes	［52］
拟南芥Arabidopsis thaliana	RST1	促进酰基辅酶A还原成醛类May act in reduction of acyl-CoAs to aldehydes	［53］
拟南芥 Arabidopsis thaliana	CYP96A15/MAH1	次级醇和酮形成Formation of secondary alcohols and ketones	［54］
拟南芥Arabidopsis thaliana	WSD1	蜡酯合成Wax ester biosynthesis	［55］
拟南芥Arabidopsis thaliana	GLN1， ECH	囊泡运输Vesicle trafficking	［56］
拟南芥Arabidopsis thaliana	LTPG1， LTPG2	蜡质转运Cuticular waxs transport	［41，57］
水稻Oryza sativa	KCR1	超长链脂肪酸合成VLCFA biosynthesis	［58］
油菜Brassica napus	CER4/FAR3	初级醇生成Formation of primary alcohols	［59］

物种 Species	基因名 Gene name	功能 Function	参考文献 Reference
拟南芥Arabidopsis thaliana	CER17	超长链酰基辅酶A的n-6去饱和 n-6 desaturation of very long chain acyl-CoAs	［4］
拟南芥Arabidopsis thaliana	LACS1， LACS2， LACS4	蜡质合成过程中重叠 Overlap in wax synthesis	［11］
拟南芥Arabidopsis thaliana	Cytb5R	CER1的辅助因子Cofactor of CER1	［19］
拟南芥Arabidopsis thaliana	WAX2/CER3	醛类合成Synthesis of aldehydes	［20］
拟南芥Arabidopsis thaliana	ABCG11/WBC11，ABCG12/CER5	蜡质转运Cuticular wax transport	［23，27］
拟南芥Arabidopsis thaliana	ACC1	丙二酰辅酶 A 底物合成 Synthesis of malonyl-CoA substrate	［29］
拟南芥Arabidopsis thaliana	KCS3	负调控蜡质合成 Negative regulation of wax biosynthesis	［30］
番茄Solanum lycopersicum	CER1-1	烷烃类合成Biosynthesis of alkanes	［34］
拟南芥Arabidopsis thaliana	KCS1	延伸C26～C30脂肪酸 Extension of C26 ~ C30 fatty acids	［43］
拟南芥Arabidopsis thaliana	CUT1， CER6， KCS5， KCS6	调控超长链脂肪酸合成 Regulation of VLCFA biosynthesis	［44］
拟南芥Arabidopsis thaliana	FATA/FATB	为蜡质合成提供饱和脂肪酸 Provide saturated fatty acids for wax biosynthesis	［45］
拟南芥Arabidopsis thaliana	CER10	超长链脂肪酸合成VLCFA biosynthesis	［46］
拟南芥Arabidopsis thaliana	PAS2/HCD	超长链脂肪酸合成VLCFA biosynthesis	［47］
拟南芥Arabidopsis thaliana	KCS20， KCS2	超长链脂肪酸扩展至C22 Extende VLCFAs to C22	［48］
拟南芥Arabidopsis thaliana	LACS1/CER8，LCAS2， LCAS4	超长链脂肪酸C20～C30 合成 Synthetase activity for VLCFAs C20-C30	［49］
拟南芥Arabidopsis thaliana	CER2， CER2-Like1/2	大于C28脂肪酸的延伸 Extension of fatty acids beyond C28	［50］
拟南芥Arabidopsis thaliana	KCS9	延伸C22～C24脂肪酸 Extension of C22～C24 fatty acids	［51］
拟南芥Arabidopsis thaliana	CER1， CER22	烷烃类合成Biosynthesis of alkanes	［52］
拟南芥Arabidopsis thaliana	RST1	促进酰基辅酶A还原成醛类May act in reduction of acyl-CoAs to aldehydes	［53］
拟南芥 Arabidopsis thaliana	CYP96A15/MAH1	次级醇和酮形成Formation of secondary alcohols and ketones	［54］
拟南芥Arabidopsis thaliana	WSD1	蜡酯合成Wax ester biosynthesis	［55］
拟南芥Arabidopsis thaliana	GLN1， ECH	囊泡运输Vesicle trafficking	［56］
拟南芥Arabidopsis thaliana	LTPG1， LTPG2	蜡质转运Cuticular waxs transport	［41，57］
水稻Oryza sativa	KCR1	超长链脂肪酸合成VLCFA biosynthesis	［58］
油菜Brassica napus	CER4/FAR3	初级醇生成Formation of primary alcohols	［59］

物种 Species	转录因子TF			受影响成分 Affected component	参考文献 Reference
物种 Species	名称 Name	类型 Type	靶基因 Target gene	受影响成分 Affected component	参考文献 Reference
油菜Brassica napus	WIN1/SHN1	AP2/EREBP	BCCP1， GPAT9， LPAT5， DGAT2， LACS2， KCS1， KCR1， CER1	C29/C31烷烃、C28/C29醇 C29/C31 alkanes， C28/C29 alcohols	［3］
小麦Triticum aestivum	MYB96-2D/5D	R2R3-MYB	CER1-6A	C27/C29/C31/C33烷烃 C27/C29/C31/C33 alkanes	［33］
拟南芥Arabidopsis thaliana	RAP2.4	AP2/DREB	KCS2， CER1	C27/C29/C31/C33烷烃 C27/C29/C31/C33 alkanes	［61］
拟南芥Arabidopsis thaliana	WRI1/3/4	AP2/ERF	PKp2， MAT， KASI， ENR1， FATA， BCCP2	二羧酸、ω-羟基脂肪酸、脂肪酸 Dicarboxylic acids， ω-hydroxy fatty acids， fatty acids	［62］
油菜Brassica napus	DEWAX1	AP1\\ERF	CER1-2， CER1	C29/C31/C33烷烃、C26脂肪酸 C29/C31/C33 alkanes， C26 fatty acid	［63］
拟南芥Arabidopsis thaliana	DEWAX2	AP2/ERF	CER1， ACLA2， LACS1， LACS2， KCS12	C29/C31烷烃、C28初级醇、C29酮 C29/C31 alkanes， C28 primary alcohol， C29 ketone	［64］
苹果Malus domestica	MYB30	R2R3-MYB	WRI1， WIN1， ACBP1， LACS2， SHINE2， SHINE3， KCS1	C29烷烃、C31醇、C29醛、C16脂肪酸、C29酮、C29/C30酯 C29 alkane， C31 alcohol， C29 aldehyde， C16 fatty acid， C29 ketone， C29/C30 esters	［66］
拟南芥Arabidopsis thaliana	MYB94， MYB96	R2R3-MYB	KCR1， KCS1， KCS2/DAISY， KCS6， CER2， CER1， CER3， WSD1	—	［67］
拟南芥Arabidopsis thaliana	MYB49	R2R3-MYB	MYB41， ABCG6， KCS	蜡质 Cuticular wax	［68］
拟南芥Arabidopsis thaliana	SPL9， SPL13	SBP-box	CER1， CER4	烷烃类、初级醇 Alkanes， primary alcohols	［6，70］
玉米Zea mays	OCL1， HDG1	HD-Zip Ⅳ	LTP， CYP78A6-like， ABCG， SEC14	C25烷烃、C24/C26/C28醇、C48酯、C28/C30醛 C25 alkane， C24/C26/C28 alcohols， C48 ester， C28/C30 aldehydes	［71］
拟南芥Arabidopsis thaliana	MYB41	R2R3-MYB	—	C22/C24脂肪醇、游离脂肪酸C22/C24 fatty alcohols， free fatty acids	［72］
拟南芥Arabidopsis thaliana	MYB96	R2R3-MYB	RD22， GH3， KCR1， SER1， KCS1， KCS2， KCS6， PAS2， CER3， ESR， WBC11， LTP	—	［73］
拟南芥Arabidopsis thaliana	MYB106，MYB16	R2R3-MYB	WIN1/SHN1， FDH， LACS2， CYP84A4， CYP77A6， KCS1， CER1， CER2， LCR， LACS2	—	［74］
水稻Oryza sativa	CFL1	ADF/cofilin	WIN1/SHN1， BDG， FDH	—	［75］

物种 Species	转录因子TF			受影响成分 Affected component	参考文献 Reference
物种 Species	名称 Name	类型 Type	靶基因 Target gene	受影响成分 Affected component	参考文献 Reference
油菜Brassica napus	WIN1/SHN1	AP2/EREBP	BCCP1， GPAT9， LPAT5， DGAT2， LACS2， KCS1， KCR1， CER1	C29/C31烷烃、C28/C29醇 C29/C31 alkanes， C28/C29 alcohols	［3］
小麦Triticum aestivum	MYB96-2D/5D	R2R3-MYB	CER1-6A	C27/C29/C31/C33烷烃 C27/C29/C31/C33 alkanes	［33］
拟南芥Arabidopsis thaliana	RAP2.4	AP2/DREB	KCS2， CER1	C27/C29/C31/C33烷烃 C27/C29/C31/C33 alkanes	［61］
拟南芥Arabidopsis thaliana	WRI1/3/4	AP2/ERF	PKp2， MAT， KASI， ENR1， FATA， BCCP2	二羧酸、ω-羟基脂肪酸、脂肪酸 Dicarboxylic acids， ω-hydroxy fatty acids， fatty acids	［62］
油菜Brassica napus	DEWAX1	AP1\\ERF	CER1-2， CER1	C29/C31/C33烷烃、C26脂肪酸 C29/C31/C33 alkanes， C26 fatty acid	［63］
拟南芥Arabidopsis thaliana	DEWAX2	AP2/ERF	CER1， ACLA2， LACS1， LACS2， KCS12	C29/C31烷烃、C28初级醇、C29酮 C29/C31 alkanes， C28 primary alcohol， C29 ketone	［64］
苹果Malus domestica	MYB30	R2R3-MYB	WRI1， WIN1， ACBP1， LACS2， SHINE2， SHINE3， KCS1	C29烷烃、C31醇、C29醛、C16脂肪酸、C29酮、C29/C30酯 C29 alkane， C31 alcohol， C29 aldehyde， C16 fatty acid， C29 ketone， C29/C30 esters	［66］
拟南芥Arabidopsis thaliana	MYB94， MYB96	R2R3-MYB	KCR1， KCS1， KCS2/DAISY， KCS6， CER2， CER1， CER3， WSD1	—	［67］
拟南芥Arabidopsis thaliana	MYB49	R2R3-MYB	MYB41， ABCG6， KCS	蜡质 Cuticular wax	［68］
拟南芥Arabidopsis thaliana	SPL9， SPL13	SBP-box	CER1， CER4	烷烃类、初级醇 Alkanes， primary alcohols	［6，70］
玉米Zea mays	OCL1， HDG1	HD-Zip Ⅳ	LTP， CYP78A6-like， ABCG， SEC14	C25烷烃、C24/C26/C28醇、C48酯、C28/C30醛 C25 alkane， C24/C26/C28 alcohols， C48 ester， C28/C30 aldehydes	［71］
拟南芥Arabidopsis thaliana	MYB41	R2R3-MYB	—	C22/C24脂肪醇、游离脂肪酸C22/C24 fatty alcohols， free fatty acids	［72］
拟南芥Arabidopsis thaliana	MYB96	R2R3-MYB	RD22， GH3， KCR1， SER1， KCS1， KCS2， KCS6， PAS2， CER3， ESR， WBC11， LTP	—	［73］
拟南芥Arabidopsis thaliana	MYB106，MYB16	R2R3-MYB	WIN1/SHN1， FDH， LACS2， CYP84A4， CYP77A6， KCS1， CER1， CER2， LCR， LACS2	—	［74］
水稻Oryza sativa	CFL1	ADF/cofilin	WIN1/SHN1， BDG， FDH	—	［75］

物种 Species	转录因子 TF			受影响成分 Affected component	参考文献 Reference
物种 Species	名称 Name	类型 Type	靶基因 Target gene	受影响成分 Affected component	参考文献 Reference
水稻Oryza sativa	WR1	ERF	KCS2， LACS1， CER3， CUT1， FDH1/2， KCS1， LACS1-2， CER1/2， FAE1-L	C16/C20/C24/C26/C30/C32 脂肪酸、C22/C32醇、C25/C27/C29/C31烷烃、C48酯C16/C20/C24/C26/C30/C32 fatty acids， C22/C32 alcohols， C25/C27/C29/C31 alkanes， C48 ester	［76］
水稻Oryza sativa	MYB60	R2R3-MYB	CER1	C29烷烃 C29 alkane	［77］
小麦Triticum aestivum	WXPL2B	AP2/ERF	KCS1	蜡质 Cuticular wax	［78］
小麦Triticum aestivum	WXPL1D	AP2/ERF	KCS1， AAT1	蜡质 Cuticular wax	［78］
小麦Triticum aestivum	MYB30	R2R3-MYB	KCS1， ECR	超长链脂肪酸、醇、醛、烷烃、酯 VLCFAs， alcohols， aldehydes， alkanes， esters	［79］
大麦Hordeum vulgare	WIN1	AP2/EREBP	KAS2， CYP86A2， CYP89A2， LACS2	—	［80］
苹果Malus domestica	SHINE2	AP2/EREBP	MYB30， MYB96， LACS2， CER1， CER3， CER6， KCS1， WIN1， DEWAX， SHINE3	烷烃、醇、醛、脂肪酸 Alkanes， alcohols， aldehydes， fatty acids	［81］
番茄Solanum lycopersicum	SHN1	ERF	—	—	［82］
菜薹Brassica rapa	SHINE3	AP2	CER2， CYP86A7， KCS2， LACS2， CER4， KCS1， CER1	脂肪酸、烷烃、醇、蜡酯 Fatty acids， alkanes， alcohols， wax esters	［83］
黄瓜Cucumis sativus	WIN1	AP2/ERF	CER1， CER1-1， CER4， KCS1， ABC	蜡酯、C29/C31烷烃 Wax esters， C29/C31 alkanes	［84］
烟草Nicotiana tabacum	MYB12a	R2R3-MYB	LOX6， LOX5， SFAR4， GDSL2	脂肪酸 Fatty acids	［85］
脐橙Citrus sinensis	MYB30	R2R3-MYB	CER1， CER4， MAH1， LTP3， WR4， MYB106	超长链脂肪酸、醇、醛、烷烃、酮 VLCFAs， alcohols， aldehydes， alkanes， ketones	［86］
洋桔梗Eustoma grandiflorum	MIXTA1	R2R3-MYB	CER3， CER6， CER10， KCS1， KCR1， CYP77A6， WIN1	蜡质 Cuticular wax	［87］