植物次生代谢的区室化研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2023-0510

摘要/Abstract

摘要：

植物次生代谢存在多层次的区室化现象，区室化对于植物的生长发育和环境胁迫响应具有重要意义。本文综述了植物次生代谢在分子水平、亚细胞水平、细胞水平和组织器官水平的区室化研究进展。次生代谢的区室化需要中间产物在不同区室间进行转运，因此转运蛋白是植物次生代谢产物区室化合成体系的重要组成部分。次生代谢区室化及其相关转运蛋白研究，丰富了植物次生代谢的理论基础，并且为天然产物合成生物学提供了新的靶点和研究策略。

关键词: 次生代谢, 区室化, 转运蛋白

Abstract:

Plant specialized metabolism（PSM）can be compartmentalized at multiple levels，which plays important roles in plant growth and development，as well as responses to environmental stress. Here we reviewed the recent advance in the compartmentalization of PSM at the molecular，subcellular，cellular and organ levels. Due to the compartmentation of PSM，the intermediates of biosynthesis pathway have to shuttle between different compartments. Therefore，the transporters are an important part of the compartmentalized biosynthesis system of specialized metabolites. The studies of the compartmentalization of PSM and its related transporters not only expand our knowledge about PSM，but also provide new targets and research strategies for the synthetic biology of natural products.

Key words: specialized metabolism, compartmentalization, transporter

关智晶, 孙超. 植物次生代谢的区室化研究进展[J]. 生物技术通报, 2024, 40(1): 1-11.

GUAN Zhi-jing, SUN Chao. Research Progress in the Compartmentalization of Plant Specialized Metabolism[J]. Biotechnology Bulletin, 2024, 40(1): 1-11.

图/表 2

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转运蛋白家族 Transporter family	次生代谢产物 Specialized metabolites	转运蛋白 Transport protein	转运蛋白序列号 GenBank	基源植物 Original species	细胞定位 Cell localization	转运方向 Transport direction	功能验证方式 Functional verification method	参考文献 Reference
ABC	小檗碱	CjMDR1（ABC B）	AB043999.1	日本黄连Coptis japonica	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[20]
		CjABCB2（ABC B）	AB674325.1	日本黄连Coptis japonica	细胞膜	向细胞内	酵母异源表达	[21]
		CjABCB3（ABC B）	AB674326.1	日本黄连Coptis japonica	细胞膜	向细胞内	酵母异源表达	[21]
	石蒜碱	LaABCB11（ABC B）	YP_009745292.1	忽地笑Lycoris aurea	细胞膜	向细胞外	酵母异源表达	[22]
	花青素	ZmMRP3（ABC C）	AAT37905.1	玉米Zea mays	液泡膜	向液泡内	酵母异源表达	[23]
	花青素	VvABCC1（ABC C）	AGC23330.1	葡萄Vitis vinifera	液泡膜	向液泡内	酵母异源表达	[24]
	花青素	AtABCC2（ABC C）	AEC09006.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[25]
	花青素	AtABCC1（ABC C）	AEE31213.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[26]
	花青素	AtABCC14（ABC C）	AEE80381.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[26]
	藏红花素	CsABCC4a（ABC C）	QEY08349.1	番红花Crocus sativus	液泡膜	向液泡内	酵母异源表达	[27]
	Sclareolide	NpPDR1（ABC G）	FE898762.1	皱叶烟草 Nicotiana plumbaginifolia	细胞膜	向细胞外	诱导蛋白表达检测底物浓度	[28]
	西松烯香紫苏醇椒二醇	NtPDR1（ABC G）	AB109388.1	烟草Nicotiana tabacum	细胞膜	—	基因过表达	[29]
	椒二醇	NbABCG1（ABC G）	LC015759.1	本氏烟草Nicotiana benthamiana	细胞膜	向细胞外	病毒诱导的基因沉默	[30]
		NbABCG2（ABC G）	LC015761	本氏烟草Nicotiana benthamiana	细胞膜	向细胞外	病毒诱导的基因沉默	[30]
	β-石竹烯	AaPDR3（ABC G）	KR153482.1	黄花蒿Artemisia annua	细胞膜	向细胞内	基因过表达、酵母异源表达	[31]
	单萜	PbABCG1（ABC G）	JQ088099.1	荧光蝴蝶兰Phalaenopsis bellina	细胞膜	向细胞外	RNA干扰基因沉默、病毒诱导的基因沉默	[32]
	挥发性有机化合物	PhABCG1（ABC G）	AFC36404.1	碧冬茄Petunia×hybrida	细胞膜	向细胞外	RNA干扰基因沉默	[33]
	长春质碱	CrTPT2（ABC G）	KC511771.1	长春花Catharanthus roseus	细胞膜	向细胞外	病毒诱导的基因沉默	[34]
	植保素Camalexin	AtABCG34（ABC G）	AEC09246.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因过表达	[35]
	香豆素	AtABCG37（ABC G）	AEE79095.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因过表达	[36]
	香豆素	NtABCG3（ABC G）	CAJ19055.1	烟草Nicotiana tabacum	细胞膜	—	病毒诱导的基因沉默、基因过表达	[37]
	苜蓿素Medicarpin	MtABCG10（ABC G）	XP_003597819	蒺藜苜蓿Medicago truncatula	细胞膜	向细胞外	基因过表达	[38]
	木质素	PgrABCG14（ABC G）	Pgr018151.1	石榴Punica granatum	细胞膜	向细胞外	基因过表达	[39]
	角质层	AtABCG32（ABC G）	AEC07906.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因敲除	[40]
	角质层	SIABCG42（ABC G）	XP_006353655.1	番茄Solanum lycopersicum	细胞膜	向细胞外	RNA干扰基因沉默	[41]
		SlABCG36（ABC G）	XP_006338166.1	番茄Solanum lycopersicum	细胞膜	向细胞外	RNA干扰基因沉默	[41]
	蜡质	ZxABCG11（ABC G）	YP_009990684.1	霸王Zygophyllum xanthoxylum	—	—	基因过表达	[42]
	脱落酸	AtABCG40（ABC G）	AEE29332.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因过表达、酵母异源表达	[43]
MATE	尼古丁	NtMATE1	AB286963.1	烟草Nicotiana tabacum	液泡膜	向液泡内	基因过表达、酵母异源表达	[44]
		NtMATE2	AB286962.1	烟草Nicotiana tabacum	液泡膜	向液泡内	基因过表达、酵母异源表达	[44]
		NtJAT1	BAG68655.1	烟草Nicotiana tabacum	液泡膜	向液泡内	酵母异源表达	[45]
		NtJAT2	BAG68656.1	烟草Nicotiana tabacum	液泡膜	向液泡内	酵母异源表达	[46]
	小檗碱	CjMATE1	BAX73926.1	日本黄连Coptis japonica	液泡膜	向液泡内	酵母异源表达	[47]
	黄酮醇	AtFFT	OAP00740.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	突变体验证	[48]
	原花青素	AtTT12	OAP05921.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[49]
	原花青素	MtMATE1	ACX37118.1	蒺藜苜蓿 Medicago truncatula	液泡膜	向液泡内	酵母异源表达	[50]
	类黄酮	MtMATE2	ADV04045.1	蒺藜苜蓿 Medicago truncatula	液泡膜	向液泡内	酵母异源表达	[51]
	原花青素	MdMATE1	GU064954	栽培苹果Malus x domestica Borkh.	液泡膜	向液泡内	突变体验证	[52]
		MdMATE2	GU064956	栽培苹果Malus x domestica Borkh.	液泡膜	向液泡内	突变体验证	[52]
	原花青素	GhTT12	AGW32085.1	陆地棉Gossypium hirsutum	液泡膜	向液泡内	突变体验证	[53]
	原花青素	FaTT12-1	AUA60209.1	草莓Fragaria×ananassa	液泡膜	向液泡内	病毒诱导的基因沉默	[54]
	花青素	VvAM1	FJ264202	葡萄Vitis vinifera	液泡膜	向液泡内	酵母异源表达	[55]
		VvAM3	FJ264203	葡萄Vitis vinifera	液泡膜	向液泡内	酵母异源表达	[55]
	异黄酮	GmMATE1	KRG94946.1	大豆Glycine max	液泡膜	向液泡内	酵母异源表达	[56]
		GmMATE2	KAG4396127.1	大豆Glycine max	液泡膜	向液泡内	酵母异源表达	[56]
	异黄酮	GmMATE4	KRH64938.1	大豆Glycine max	液泡膜	向液泡内	酵母异源表达	[57]
	CCoumaroylagmatine	AtDTX18	AEE76776.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	突变体验证	[58]
	黄酮醇	NtMATE21	XP_016475205.1	烟草Nicotiana tabacum	细胞膜	向细胞外	突变体验证	[59]
		NtMATE22	XP_016477351.1	烟草Nicotiana tabacum	细胞膜	向细胞外	突变体验证	[59]
	染料木素	LaMATE2	KY464927	白羽扇豆Lupinus albus	细胞膜	向细胞外	病毒诱导的基因沉默	[60]
	苦味素	CsMATE1	AXN55888.1	黄瓜Cucumis sativus	液泡膜	向液泡内	基因过表达	[61]
	水杨酸	EDS5	ABZ03276.1	拟南芥Arabidopsis thaliana	叶绿体被膜	向叶绿体外	酵母异源表达	[62]
PUP	尼古丁	NUP1	ADP30799.1	烟草Nicotiana tabacum	细胞膜	向细胞内	酵母异源表达	[63]
	苄基异喹啉生物碱	BUP1	QBG64391.1	罂粟Papaver somniferum	细胞膜	向细胞内	酵母异源表达	[64]
	咖啡因	CsPUP1	TEA003596	茶树Camellia sinensis	细胞膜	向细胞内	基因过表达	[65]
		CsPUP3.1	TEA029223	茶树Camellia sinensis	细胞膜	向细胞内	基因过表达	[65]
		CsPUP10.1	TEA023430	茶树Camellia sinensis	细胞膜	向细胞内	基因过表达	[65]
NPF	硫代葡萄糖苷	AtNPF2.9（AtGTR3）	Q9M9V7.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[66]
	硫代葡萄糖苷	AtNPF2.10（AtGTR1）	Q944G5.3	拟南芥Arabidopsis thaliana	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[67]
	硫代葡萄糖苷	AtNPF2.11（AtGTR2）	BAH19623.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[67]
	黄酮醇苷	AtFST1（NPF2.8）	Q3E8X3.2	拟南芥Arabidopsis thaliana	细胞膜	—	大肠杆菌异源表达	[68]
	长春碱和长春新碱	CrNPF2.9	AQM73449.1	长春花Catharanthus roseus	液泡膜	向液泡外	病毒诱导的基因沉默	[69]
	长春碱和长春新碱	CrNPF2.4	ALE20039.1	长春花Catharanthus roseus	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[70]
	长春碱和长春新碱	CrNPF 2.5	ALE20040.1	长春花Catharanthus roseus	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[70]
	长春碱和长春新碱	CrNPF 2.6	ALE20041.1	长春花Catharanthus roseus	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[70]
	番茄碱	SlNPF1.5	OP765903.1	番茄Solanum lycopersicum	液泡膜	向液泡外	爪蟾卵母细胞异源表达	[71]

转运蛋白家族 Transporter family	次生代谢产物 Specialized metabolites	转运蛋白 Transport protein	转运蛋白序列号 GenBank	基源植物 Original species	细胞定位 Cell localization	转运方向 Transport direction	功能验证方式 Functional verification method	参考文献 Reference
ABC	小檗碱	CjMDR1（ABC B）	AB043999.1	日本黄连Coptis japonica	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[20]
		CjABCB2（ABC B）	AB674325.1	日本黄连Coptis japonica	细胞膜	向细胞内	酵母异源表达	[21]
		CjABCB3（ABC B）	AB674326.1	日本黄连Coptis japonica	细胞膜	向细胞内	酵母异源表达	[21]
	石蒜碱	LaABCB11（ABC B）	YP_009745292.1	忽地笑Lycoris aurea	细胞膜	向细胞外	酵母异源表达	[22]
	花青素	ZmMRP3（ABC C）	AAT37905.1	玉米Zea mays	液泡膜	向液泡内	酵母异源表达	[23]
	花青素	VvABCC1（ABC C）	AGC23330.1	葡萄Vitis vinifera	液泡膜	向液泡内	酵母异源表达	[24]
	花青素	AtABCC2（ABC C）	AEC09006.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[25]
	花青素	AtABCC1（ABC C）	AEE31213.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[26]
	花青素	AtABCC14（ABC C）	AEE80381.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[26]
	藏红花素	CsABCC4a（ABC C）	QEY08349.1	番红花Crocus sativus	液泡膜	向液泡内	酵母异源表达	[27]
	Sclareolide	NpPDR1（ABC G）	FE898762.1	皱叶烟草 Nicotiana plumbaginifolia	细胞膜	向细胞外	诱导蛋白表达检测底物浓度	[28]
	西松烯香紫苏醇椒二醇	NtPDR1（ABC G）	AB109388.1	烟草Nicotiana tabacum	细胞膜	—	基因过表达	[29]
	椒二醇	NbABCG1（ABC G）	LC015759.1	本氏烟草Nicotiana benthamiana	细胞膜	向细胞外	病毒诱导的基因沉默	[30]
		NbABCG2（ABC G）	LC015761	本氏烟草Nicotiana benthamiana	细胞膜	向细胞外	病毒诱导的基因沉默	[30]
	β-石竹烯	AaPDR3（ABC G）	KR153482.1	黄花蒿Artemisia annua	细胞膜	向细胞内	基因过表达、酵母异源表达	[31]
	单萜	PbABCG1（ABC G）	JQ088099.1	荧光蝴蝶兰Phalaenopsis bellina	细胞膜	向细胞外	RNA干扰基因沉默、病毒诱导的基因沉默	[32]
	挥发性有机化合物	PhABCG1（ABC G）	AFC36404.1	碧冬茄Petunia×hybrida	细胞膜	向细胞外	RNA干扰基因沉默	[33]
	长春质碱	CrTPT2（ABC G）	KC511771.1	长春花Catharanthus roseus	细胞膜	向细胞外	病毒诱导的基因沉默	[34]
	植保素Camalexin	AtABCG34（ABC G）	AEC09246.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因过表达	[35]
	香豆素	AtABCG37（ABC G）	AEE79095.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因过表达	[36]
	香豆素	NtABCG3（ABC G）	CAJ19055.1	烟草Nicotiana tabacum	细胞膜	—	病毒诱导的基因沉默、基因过表达	[37]
	苜蓿素Medicarpin	MtABCG10（ABC G）	XP_003597819	蒺藜苜蓿Medicago truncatula	细胞膜	向细胞外	基因过表达	[38]
	木质素	PgrABCG14（ABC G）	Pgr018151.1	石榴Punica granatum	细胞膜	向细胞外	基因过表达	[39]
	角质层	AtABCG32（ABC G）	AEC07906.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因敲除	[40]
	角质层	SIABCG42（ABC G）	XP_006353655.1	番茄Solanum lycopersicum	细胞膜	向细胞外	RNA干扰基因沉默	[41]
		SlABCG36（ABC G）	XP_006338166.1	番茄Solanum lycopersicum	细胞膜	向细胞外	RNA干扰基因沉默	[41]
	蜡质	ZxABCG11（ABC G）	YP_009990684.1	霸王Zygophyllum xanthoxylum	—	—	基因过表达	[42]
	脱落酸	AtABCG40（ABC G）	AEE29332.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	基因过表达、酵母异源表达	[43]
MATE	尼古丁	NtMATE1	AB286963.1	烟草Nicotiana tabacum	液泡膜	向液泡内	基因过表达、酵母异源表达	[44]
		NtMATE2	AB286962.1	烟草Nicotiana tabacum	液泡膜	向液泡内	基因过表达、酵母异源表达	[44]
		NtJAT1	BAG68655.1	烟草Nicotiana tabacum	液泡膜	向液泡内	酵母异源表达	[45]
		NtJAT2	BAG68656.1	烟草Nicotiana tabacum	液泡膜	向液泡内	酵母异源表达	[46]
	小檗碱	CjMATE1	BAX73926.1	日本黄连Coptis japonica	液泡膜	向液泡内	酵母异源表达	[47]
	黄酮醇	AtFFT	OAP00740.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	突变体验证	[48]
	原花青素	AtTT12	OAP05921.1	拟南芥Arabidopsis thaliana	液泡膜	向液泡内	酵母异源表达	[49]
	原花青素	MtMATE1	ACX37118.1	蒺藜苜蓿 Medicago truncatula	液泡膜	向液泡内	酵母异源表达	[50]
	类黄酮	MtMATE2	ADV04045.1	蒺藜苜蓿 Medicago truncatula	液泡膜	向液泡内	酵母异源表达	[51]
	原花青素	MdMATE1	GU064954	栽培苹果Malus x domestica Borkh.	液泡膜	向液泡内	突变体验证	[52]
		MdMATE2	GU064956	栽培苹果Malus x domestica Borkh.	液泡膜	向液泡内	突变体验证	[52]
	原花青素	GhTT12	AGW32085.1	陆地棉Gossypium hirsutum	液泡膜	向液泡内	突变体验证	[53]
	原花青素	FaTT12-1	AUA60209.1	草莓Fragaria×ananassa	液泡膜	向液泡内	病毒诱导的基因沉默	[54]
	花青素	VvAM1	FJ264202	葡萄Vitis vinifera	液泡膜	向液泡内	酵母异源表达	[55]
		VvAM3	FJ264203	葡萄Vitis vinifera	液泡膜	向液泡内	酵母异源表达	[55]
	异黄酮	GmMATE1	KRG94946.1	大豆Glycine max	液泡膜	向液泡内	酵母异源表达	[56]
		GmMATE2	KAG4396127.1	大豆Glycine max	液泡膜	向液泡内	酵母异源表达	[56]
	异黄酮	GmMATE4	KRH64938.1	大豆Glycine max	液泡膜	向液泡内	酵母异源表达	[57]
	CCoumaroylagmatine	AtDTX18	AEE76776.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞外	突变体验证	[58]
	黄酮醇	NtMATE21	XP_016475205.1	烟草Nicotiana tabacum	细胞膜	向细胞外	突变体验证	[59]
		NtMATE22	XP_016477351.1	烟草Nicotiana tabacum	细胞膜	向细胞外	突变体验证	[59]
	染料木素	LaMATE2	KY464927	白羽扇豆Lupinus albus	细胞膜	向细胞外	病毒诱导的基因沉默	[60]
	苦味素	CsMATE1	AXN55888.1	黄瓜Cucumis sativus	液泡膜	向液泡内	基因过表达	[61]
	水杨酸	EDS5	ABZ03276.1	拟南芥Arabidopsis thaliana	叶绿体被膜	向叶绿体外	酵母异源表达	[62]
PUP	尼古丁	NUP1	ADP30799.1	烟草Nicotiana tabacum	细胞膜	向细胞内	酵母异源表达	[63]
	苄基异喹啉生物碱	BUP1	QBG64391.1	罂粟Papaver somniferum	细胞膜	向细胞内	酵母异源表达	[64]
	咖啡因	CsPUP1	TEA003596	茶树Camellia sinensis	细胞膜	向细胞内	基因过表达	[65]
		CsPUP3.1	TEA029223	茶树Camellia sinensis	细胞膜	向细胞内	基因过表达	[65]
		CsPUP10.1	TEA023430	茶树Camellia sinensis	细胞膜	向细胞内	基因过表达	[65]
NPF	硫代葡萄糖苷	AtNPF2.9（AtGTR3）	Q9M9V7.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[66]
	硫代葡萄糖苷	AtNPF2.10（AtGTR1）	Q944G5.3	拟南芥Arabidopsis thaliana	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[67]
	硫代葡萄糖苷	AtNPF2.11（AtGTR2）	BAH19623.1	拟南芥Arabidopsis thaliana	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[67]
	黄酮醇苷	AtFST1（NPF2.8）	Q3E8X3.2	拟南芥Arabidopsis thaliana	细胞膜	—	大肠杆菌异源表达	[68]
	长春碱和长春新碱	CrNPF2.9	AQM73449.1	长春花Catharanthus roseus	液泡膜	向液泡外	病毒诱导的基因沉默	[69]
	长春碱和长春新碱	CrNPF2.4	ALE20039.1	长春花Catharanthus roseus	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[70]
	长春碱和长春新碱	CrNPF 2.5	ALE20040.1	长春花Catharanthus roseus	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[70]
	长春碱和长春新碱	CrNPF 2.6	ALE20041.1	长春花Catharanthus roseus	细胞膜	向细胞内	爪蟾卵母细胞异源表达	[70]
	番茄碱	SlNPF1.5	OP765903.1	番茄Solanum lycopersicum	液泡膜	向液泡外	爪蟾卵母细胞异源表达	[71]