HD-ZIP转录因子响应病菌与非生物胁迫研究进展

刘娇; 帅鹏

doi:10.19303/j.issn.1008-0384.2021.01.015

HD-ZIP转录因子响应病菌与非生物胁迫研究进展

doi: 10.19303/j.issn.1008-0384.2021.01.015

刘娇,
帅鹏^,

福建农林大学林学院，福建　福州　350002

基金项目: 国家自然科学基金项目（31971626）；国家自然基金青年科学基金项目（31600481）

详细信息

作者简介:
刘娇（1997−），女，硕士研究生，主要从事植物分子机制研究（E-mail: Lioujiao-161726084@qq.com）

通讯作者:
帅鹏（1988−），男，硕士生导师，主要从事植物分子机制研究（E-mail: grooot@qq.com）

中图分类号: Q 945
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- 被引次数: 0
出版历程
- 收稿日期: 2020-08-16
- 修回日期: 2020-10-21
- 网络出版日期: 2020-11-13
- 刊出日期: 2021-01-31

Research Progress on HD-ZIP Transcription Factors in Response to Pathogenic or Abiotic Stresses

LIU Jiao,
SHUAI Peng^,

Fujian Agriculture and Forestry University, Fuzhou, Fujian　350002, China

摘要

摘要: 逆境条件通常影响植物生长发育，间接或直接导致作物减产甚至植物死亡。HD-ZIP转录因子则参与植物对不利环境条件的响应。Homeodomain-Leucine Zipper（HD-ZIP）转录因子是植物中特有的一类转录因子，属于同源异形盒（homeobox, HB）蛋白家族，由高度保守的同源异形结构域（Homeodomain）和亮氨酸拉链结构域（ZIP）紧密连接而成。通过LZ结构域介导的蛋白二聚体的形成使HD结构域与靶DNA结合，调控靶基因的表达。HD-ZIP转录因子不仅对植物生长发育发挥重要调控作用，并且对逆境抵抗中起关键作用。本文基于近年来HD-ZIP转录因子的最新研究成果，着重归纳HD-ZIP四个亚家族（Ⅰ-Ⅳ）对不同病菌和非生物胁迫例如干旱、高盐、极端温度、弱光、机械损伤、重金属胁迫做出的响应机制，以期揭示HD-ZIP转录因子如何通过整合激素和环境信号来改良植物生长特性的内在分子机制，从而为提升植物抗逆性奠定基础。
- HD-ZIP /
- 病菌胁迫 /
- 非生物胁迫 /
- 植物抗逆性
Abstract: Adverse external conditions commonly affect plant growth and development which directly or indirectly cause decline on crop yield and even death of the plants. The homeodomain-leucine zipper (HD-ZIP) transcription factors have been known to involve in stress responses of plants. Belonging to the homeobox (HB) protein family, the factors are unique to plants and tightly connected by the highly conserved HD and ZIP. The formation of protein dimers mediated by the LZ domain allows HD to bind to the target DNA and regulate the expression of the target gene. The HD-ZIP transcription factors not only play an important role in regulating plant growth and development but also in responding to external stresses. This article focuses on the published reports of recent studies concerning the roles of the 4 subfamilies Ⅰ-Ⅳ of the HD-ZIP transcription factors in response to pathogenic attacks and/or abiotic stresses such as drought, salt, extreme temperature, wounding, low R/FR light, and heavy metals. Through the internal molecular response mechanisms initiated by HD-ZIP, a plant could ward off the imposed adversities. With an in-depth understanding of the functions, means to improve the growth and stress resistance of plants could be realized.
- HD-ZIP /
- pathogenic stress /
- abiotic stress /
- stress resistance of plants

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图 1 HD-ZIP家族的分类结构示意图^[15]

Figure 1. Schematic diagram of classified HD-ZIP family^[15]

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图 2 HD-ZIP干旱胁迫调控通路

Figure 2. Regulatory route of HD-ZIP in response to drought stress

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图 3 HD-ZIP盐胁迫调控通路

Figure 3. Regulatory route of HD-ZIP in response to salt stress

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图 4 HD-ZIP避光反应调控通路

注：R/FR光的变化导致R光吸收模式（Pr）和FR光吸收模式（Pfr）之间的平衡向Pr转移，从而导致植物色素PhyB、PhyD、PhyE失活。植物色素失活有利于避光反应正调节因子PIFs的转录。PIF进而激活HD-ZIP Ⅱ和HFR1/SICS1基因的转录。HFR1/SICS1则会抑制PIF和ATHB2、ATHB4的活性。然而PhyB、PhyD、PhyE以及HD-ZIP Ⅲ亚家族成员可激活ATHB2、ATHB4的转录活性从而正调控避光反应。

Figure 4. Regulatory route of HD-ZIP in response to shading

Note: Changes in R/FR light cause equilibrium sift from FR light-absorbing photo-convertible isoform (Pfr) to R light-absorbing photo-convertible isoform (Pr) resulting in deactivations of phyB, phyD, and phyE. This, in turn, results in an enhanced stability and/or activity of several phytochrome-interacting transcription factors (PIFs). PIFs, within a brief moment, activate the transcription of HD-ZIPs II and HFR1/SICS1 genes. HFR1/SICS1 can inhibit activities of PIF, ATHB2, and ATHB4. But PhyB, PhyD, PhyE, and HD-ZIP Ⅲ subfamily members can activate the transcriptional activities of ATHB2 and ATHB4 to positively regulate the response to light-shielding.

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表 1 HD-ZIP在干旱胁迫下的不同表达模式

Table 1. Differential expressions of HD-ZIP transcription factors under drought stress

	植物物种 Plant species	HD-ZIP转录因子 HD-ZIP TFs	亚家族 Subfamily	表达模式 Express patterns
单子叶植物 monocotyledon	玉米 Zea mays L.	ZmHDZ4	Ⅰ	1 h时显著上调 up-regulated Significantly at 1 h
		Zmhdz10	Ⅰ	上调，12时达到峰值 up-regulated,peaked at 12 h
	水稻 Oryza sativa L.	OsHOX22	Ⅰ	显著上调 up-regulated Significantly
	水稻 Oryza sativa L.	OsHOX24	Ⅰ	显著上调 up-regulated Significantly
	小麦 Triticum aestivum	TaHDZipI-3	Ⅰ	微弱上调 Slightly up-regulated
	小麦 Triticum aestivum	TaHDZipI-4	Ⅰ	较为显著上调 up-regulated
		TaHDZipI-5	Ⅰ	显著上调 up-regulated Significantly
双子叶植物 Dicotyledon	拟南芥 Arabidopsis thaliana	AtHB13	Ⅰ	上调，8天时达到5倍峰值 up-regulated，fivefold after 8 days
	拟南芥 Arabidopsis thaliana	AtHB7AtHB12	Ⅰ	显著上调 up-regulated Significantly
		HDG11	Ⅳ	上调 up-regulated
	向日葵 Helianthus annuus L.	HaHB4	Ⅰ	上调 up-regulated
	鹰嘴豆 Cicer arietinum	CaHDZ12	Ⅰ	根中48 h上调显著 up-regulated Significantly in roots after 48 h
	番茄 Solanum lycopersicum	SIHB2	Ⅰ	2 h后持续上调，保持高水平 Continue to up-regulatedand keep a high level after 2 h

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表 2 HD-ZIP转录因子在盐胁迫下不同表达模式

Table 2. Differential expressions of HD-ZIP transcription factors under salt stress

项目 Items	植物物种 Plant species	HD-ZIP TFs	亚家族 Subfamily	表达模式 Express patterns
单子叶植物 monocotyledon	玉米 Zea mays L.	ZmHDZ10	Ⅰ	上调，12 h达到峰值 up-regulated,peaked at 12 h
单子叶植物 monocotyledon	玉米 Zea mays L.	ZmHDZ1	Ⅰ	1 h时显著上调 up-regulated Significantly at 1 h
双子叶植物 Dicotyledon	拟南芥 Arabidopsis thaliana	ATHB17	Ⅰ	在叶中显著诱导 Induced Significantly in leaves
	拟南芥 Arabidopsis thaliana	HDG11	Ⅳ	上调 up-regulated
	毛果杨 Populus trichocarpa	PtrHox11		上调 up-regulated
	大豆 Glycine max	Gshdz4	Ⅰ	在根中显著上调 up-regulated Significantly in roots
	鹰嘴豆 Cicerarietinum	CaHDZ12	Ⅰ	在72 h、96 h显著上调 up-regulated Significantly at 72 h,96 h
	番茄 Solanum lycopersicum	SIHB2	Ⅰ	在2 h时显著上调 up-regulated Significantly at 2 h
	麻风树 Jatropha curcas L.	JcHDZ16	Ⅰ	所有时间点均被下调 Down-regulated at all pont

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表 3 HD-ZIP转录因子在高温与低温胁迫中的不同表达模式

Table 3. Differential expressions of HD-ZIP transcription factors under high- and low-temp stresses

植物物种 Plant species	HD-ZIP转录因子 HD-ZIP TFs	亚家族 Subfamily	温度胁迫 Temperature stress	表达模式 Expression Patterns
黄瓜 Cucumis sativus L.	CsHDZ02，33	Ⅰ	高温低温	显著上调 up-regulated Significantly
	CsHDZ08，22	Ⅱ
	CsHDZ11，37	Ⅳ
土豆 Solanum tuberosum L.	StHOX20	Ⅱ		在根与叶中显著上调 up-regulated Significantly in leaves and roots
小麦 Triticum aestivum L.	TaHDZipI-2TaHDZipI-3	Ⅰ	低温	下调 down-regulated
小麦 Triticum aestivum L.	TaHDZipI-4TaHDZipI-5	Ⅰ		上调 up-regulated
番茄 Solanum lycopersicum	SIHZ17	Ⅰ		下调，且没有反弹 down-regulated without any rebound
黑麦草 Lolium perenne L.	LpHOX6，8，24	Ⅰ	高温	在叶片和根中上调 up-regulated in leaves and roots
黑麦草 Lolium perenne L.	LpHOX21	Ⅰ		在根和叶片中下调 down-regulated in leaves and roots
向日葵 Helianthus annuus L.	HaHB4	Ⅰ		上调 up-regulated

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