WRKY对糖调控园艺作物冷适应的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2021-0691

摘要/Abstract

摘要：

植物的糖既能参与细胞的碳和能量代谢,又能作为信号分子促进植株生长发育并参与调控植物对逆境胁迫的响应。目前诸多研究表明抗寒锻炼中可溶性糖的积累有助于保护植物抵御冻害,黄瓜等园艺作物的抗冷性在外源施糖后提高,但具体机制尚未明确。糖能够促进植株通过表观形态、生理生化及分子水平等方面对非生物逆境胁迫响应、调控植株的抗性。糖代谢过程与脱落酸（ABA）等植物激素的调控密切相关。WRKY作为ABA相关逆境应答信号通路中的核心转录因子,同是糖代谢调控机制中的重要因子,或许在响应冷胁迫的相关代谢机制中参与发挥着重要作用。综述了糖对园艺作物冷适应的响应及调控的分子机理,并分析讨论了WRKY家族对园艺植物糖调控冷适应效应响应的机理。

关键词: 低温, 转录因子, WRKY, 外源糖

Abstract:

Sugar in plant can not only promote the cellular carbon metabolism and energy metabolism,also act as a signal molecule to promote plant growth and participate in the regulation of plant response to adversity stress. Currently,a number of studies reveal that the accumulation of soluble sugar is conducive to protecting plants from freezing injury in cold hardening,it has been shown that the cold resistance of horticultural crops such as cucumber has been improved under exogenous sugar application,however,the specific mechanisms are not clear yet. Sugar can promote plants’ responses to abiotic stress through different regulatory mechanisms,such as physiology and biochemistry,and different molecular regulatory mechanisms. WRKY,as a core transcription factor in the ABA-related adversity response signaling pathway,is also an important factor in the sugar metabolism regulation mechanisms,and may play an important role in the related metabolic mechanisms in response to cold stress. Here the response and regulatory mechanisms of cold acclimation in horticultural crops by sugar are reviewed,and the role of WRKY family in the process is analyzed and discussed.

Key words: cold stress, transcription factor, WRKY, exogenous sugar

潘英杰, 张颖, 武杞蔓, 李正青. WRKY对糖调控园艺作物冷适应的研究进展[J]. 生物技术通报, 2022, 38(3): 203-212.

PAN Ying-jie, ZHANG Ying, WU Qi-man, LI Zheng-qing. A Review of WRKY Mediated Regulation of Sugar for Cold Acclimation in Horticultural Crops[J]. Biotechnology Bulletin, 2022, 38(3): 203-212.

图/表 2

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逆境 Stress	物种 Species	外源糖 Exogenous sugar	变量Variable				参考文献 Reference
逆境 Stress	物种 Species	外源糖 Exogenous sugar	品质 Quality	生理、生化 Physiology,biochemistry	激素 Hormone	分子机理 Molecular mechanisms	参考文献 Reference
无 None	苹果 Apple	ALA	果实着色、外观品质↑ Fruit coloring and appearance quality 果皮花青素含量↑ Anthocyanin content in pericarp				[12]
	草莓 Strawberry	蔗糖 Sucrose	花青素和类胡萝卜素含量↑ Anthocyanin and carotenoid content	抗坏血酸含量↑ Ascorbic acid content			[13]
	蓝莓 Blueberry	蔗糖 Sucrose	可溶性总糖、花色苷含量↑ Total soluble sugar and anthocyanin 可滴定酸含量↓Titratable acid	黄酮和抗氧化能力↓ Flavonoids and antioxidant capacity			[14]
	苹果 Apple	葡萄糖 Glucose	短枝顶芽长度、宽度及质量↑ Length,width and quality of terminal bud of short branch 淀粉↑ Amylopectin	酸性与中性蔗糖转化酶活性↓ Acid and neutral sucrose invertase activity 蔗糖合酶分解方向酶活性↓ Enzyme activity in the direction of sucrose synthase decomposition		顶芽糖代谢相关基因及Genes related to glucose metabolism in apical buds 2个MdTPS基因响应 2 MdTPS genes response 成花关键基因 ↑Key genes for flowering	[15]
	茄子Eggplant	葡萄糖 Glucose	叶片变绿The leaves turn green 节间距↓Internode distance 可溶性糖↑Soluble sugars	抗氧化酶活性↑Antioxidant enzyme activity 叶片质膜相对透性↓Relative permeability of plasma membrane of leaves			[16]
冷胁迫 Cold stress	番茄 Tomato	ALA		叶绿素合成↑ Chlorophyll synthesis 膜脂过氧化↓ Membrane lipid peroxidation	内源H₂O₂、JA、NO参与 Internal source H₂O₂,JA,and NO	RBOH1、GSTU43表达↑ RBOH1 and GSTU43 expression	[17]
	茶树 Tea tree	ALA	茶叶品质↑ Tea quality	光合能力↑ Photosynthetic capacity			[18]
	玉米 Corn	海藻糖 Trehalose	根表面积、根长、 Root surface area and root length 鲜干重↑Fresh and dry weight	渗透物质的积累↑Accumulation of osmotic substances 膜脂过氧化↓Membrane lipid peroxidation			[19]
	黄瓜 Cucumber	海藻糖 Trehalose		叶片膜脂过氧化↓Membrane lipid peroxidation of leaves 叶绿素含量↑Chlorophyll content			[1]

逆境 Stress	物种 Species	外源糖 Exogenous sugar	变量Variable				参考文献 Reference
逆境 Stress	物种 Species	外源糖 Exogenous sugar	品质 Quality	生理、生化 Physiology,biochemistry	激素 Hormone	分子机理 Molecular mechanisms	参考文献 Reference
无 None	苹果 Apple	ALA	果实着色、外观品质↑ Fruit coloring and appearance quality 果皮花青素含量↑ Anthocyanin content in pericarp				[12]
	草莓 Strawberry	蔗糖 Sucrose	花青素和类胡萝卜素含量↑ Anthocyanin and carotenoid content	抗坏血酸含量↑ Ascorbic acid content			[13]
	蓝莓 Blueberry	蔗糖 Sucrose	可溶性总糖、花色苷含量↑ Total soluble sugar and anthocyanin 可滴定酸含量↓Titratable acid	黄酮和抗氧化能力↓ Flavonoids and antioxidant capacity			[14]
	苹果 Apple	葡萄糖 Glucose	短枝顶芽长度、宽度及质量↑ Length,width and quality of terminal bud of short branch 淀粉↑ Amylopectin	酸性与中性蔗糖转化酶活性↓ Acid and neutral sucrose invertase activity 蔗糖合酶分解方向酶活性↓ Enzyme activity in the direction of sucrose synthase decomposition		顶芽糖代谢相关基因及Genes related to glucose metabolism in apical buds 2个MdTPS基因响应 2 MdTPS genes response 成花关键基因 ↑Key genes for flowering	[15]
	茄子Eggplant	葡萄糖 Glucose	叶片变绿The leaves turn green 节间距↓Internode distance 可溶性糖↑Soluble sugars	抗氧化酶活性↑Antioxidant enzyme activity 叶片质膜相对透性↓Relative permeability of plasma membrane of leaves			[16]
冷胁迫 Cold stress	番茄 Tomato	ALA		叶绿素合成↑ Chlorophyll synthesis 膜脂过氧化↓ Membrane lipid peroxidation	内源H₂O₂、JA、NO参与 Internal source H₂O₂,JA,and NO	RBOH1、GSTU43表达↑ RBOH1 and GSTU43 expression	[17]
	茶树 Tea tree	ALA	茶叶品质↑ Tea quality	光合能力↑ Photosynthetic capacity			[18]
	玉米 Corn	海藻糖 Trehalose	根表面积、根长、 Root surface area and root length 鲜干重↑Fresh and dry weight	渗透物质的积累↑Accumulation of osmotic substances 膜脂过氧化↓Membrane lipid peroxidation			[19]
	黄瓜 Cucumber	海藻糖 Trehalose		叶片膜脂过氧化↓Membrane lipid peroxidation of leaves 叶绿素含量↑Chlorophyll content			[1]

物种 Species	数目 Number	已鉴定功能 Identified function			响应机制 Response mechanism	参考文献 Reference
物种 Species	数目 Number	生长发育 Growth	病原体响应 Response to pathogen	低温响应 Response to low temperature	响应机制 Response mechanism	参考文献 Reference
番茄 Tomato	86	多数I类如SlWRKY2、SlWRKY63、 SlWRKY49	SlWPKY78、SlWPKY14、SlWPKY58、SlWPKY49、SlWPKY21、SlWPKY72、SlWPKY68	SlWRKY2、SlWRKY6、SlWRKY9、SlWRKY11、SlWRKY21、SlWRKY24、SlWRKY35、SlWRKY43、SlWRKY50、SlWRKY59、SlWRKY69、SlWRKY23、SlWRKY24、SlWRKY35、SlWRKY43、SlWRKY49、SlWRKY59、SlWRKY62、SlWRKY1、SlWRKY8、SlWRKY61、SlWRKY79等	SlWRKY复合体发挥功能 SlWRKY-complex plays its function ICE-CBF-COR途径;与其他转录因子互作 ICE-CBF-COR pathways;interaction with other transcription factors	[40,43]
黄瓜Cucumber	57	CsWRKY25、CsWRKY32、CsWRKY50、 CsWRKY52、 CsWRKY59		CsWRKY18、CsWRKY21、CsWRKY25、CsWRKY40、CsWRKY57 显著上调表达（strong induction of gene expression） CsWRKY2、CsWRKY4、CsWRKY23、CsWRKY33、CsWRKY36、CsWRKY46上调表达（moderate induction of gene expression） CsWRKY28、CsWRKY53、CsWRKY55下调表达（reduction of gene expression）	CsWRKY2、CsWRKY32与MAPK7互作 CsWRKY2 and CsWRKY32 interaction with MAPK7 CsWRKY46与ABI5启动子中的W-box相互作用 CsWRKY46 and ABI5 interaction with W-box	[41,44-45]
草莓 Strawberry	62			FvWRKY27、FvWRKY41、FvWRKY56、FvWRKY62等9个上调表达 32个下调表达		[46]
西瓜 Watermelon	57	56个ClWRKY		7个上调表达,Cla018059（ClWRKY55）和 Cla021170（ClWRKY20）;12个下调表达	抗氧化能力↑ Antioxidant capacity	[47]
葡萄 Grape	59			VvWRKY55、VvWRKY14、VvWRKY12、VvWRKY52、VvWRKY28、VvWRKY43、VvWRKY49、VvWRKY37、VvWRKY08、VvWRKY58、VvWRKY33、VvWRKY39、VvWRKY19、VvWRKY47、VvWRKY44、VvWRKY51、VvWRKY45、VvWRKY31、VvWRKY06、VvWRKY34、VvWRKY26、VvWRKY53上调表达 VvWRKY11、VvWPKY16、VvWPKY22、VvWPKY29、VvWPKY41下调表达	3个VvWRKY（VvWRKY28、42和55）可能参与ABA依赖的信号通路 3 VvWRKY（VvWRKY28,42 and 55 participate in ABA-dependent signaling pathways 另外12个VvWRKY可能参与ABA不依赖的信号通路 Other 12 VvWRKYs participate in ABA-independent signaling pathways	[48]

物种 Species	数目 Number	已鉴定功能 Identified function			响应机制 Response mechanism	参考文献 Reference
物种 Species	数目 Number	生长发育 Growth	病原体响应 Response to pathogen	低温响应 Response to low temperature	响应机制 Response mechanism	参考文献 Reference
番茄 Tomato	86	多数I类如SlWRKY2、SlWRKY63、 SlWRKY49	SlWPKY78、SlWPKY14、SlWPKY58、SlWPKY49、SlWPKY21、SlWPKY72、SlWPKY68	SlWRKY2、SlWRKY6、SlWRKY9、SlWRKY11、SlWRKY21、SlWRKY24、SlWRKY35、SlWRKY43、SlWRKY50、SlWRKY59、SlWRKY69、SlWRKY23、SlWRKY24、SlWRKY35、SlWRKY43、SlWRKY49、SlWRKY59、SlWRKY62、SlWRKY1、SlWRKY8、SlWRKY61、SlWRKY79等	SlWRKY复合体发挥功能 SlWRKY-complex plays its function ICE-CBF-COR途径;与其他转录因子互作 ICE-CBF-COR pathways;interaction with other transcription factors	[40,43]
黄瓜Cucumber	57	CsWRKY25、CsWRKY32、CsWRKY50、 CsWRKY52、 CsWRKY59		CsWRKY18、CsWRKY21、CsWRKY25、CsWRKY40、CsWRKY57 显著上调表达（strong induction of gene expression） CsWRKY2、CsWRKY4、CsWRKY23、CsWRKY33、CsWRKY36、CsWRKY46上调表达（moderate induction of gene expression） CsWRKY28、CsWRKY53、CsWRKY55下调表达（reduction of gene expression）	CsWRKY2、CsWRKY32与MAPK7互作 CsWRKY2 and CsWRKY32 interaction with MAPK7 CsWRKY46与ABI5启动子中的W-box相互作用 CsWRKY46 and ABI5 interaction with W-box	[41,44-45]
草莓 Strawberry	62			FvWRKY27、FvWRKY41、FvWRKY56、FvWRKY62等9个上调表达 32个下调表达		[46]
西瓜 Watermelon	57	56个ClWRKY		7个上调表达,Cla018059（ClWRKY55）和 Cla021170（ClWRKY20）;12个下调表达	抗氧化能力↑ Antioxidant capacity	[47]
葡萄 Grape	59			VvWRKY55、VvWRKY14、VvWRKY12、VvWRKY52、VvWRKY28、VvWRKY43、VvWRKY49、VvWRKY37、VvWRKY08、VvWRKY58、VvWRKY33、VvWRKY39、VvWRKY19、VvWRKY47、VvWRKY44、VvWRKY51、VvWRKY45、VvWRKY31、VvWRKY06、VvWRKY34、VvWRKY26、VvWRKY53上调表达 VvWRKY11、VvWPKY16、VvWPKY22、VvWPKY29、VvWPKY41下调表达	3个VvWRKY（VvWRKY28、42和55）可能参与ABA依赖的信号通路 3 VvWRKY（VvWRKY28,42 and 55 participate in ABA-dependent signaling pathways 另外12个VvWRKY可能参与ABA不依赖的信号通路 Other 12 VvWRKYs participate in ABA-independent signaling pathways	[48]