拟南芥COPT家族蛋白研究进展

doi:10.11983/CBB19045

摘要/Abstract

摘要： 铜(Cu)是植物必需的微量元素, 作为多种酶的辅因子参与许多植物生理生化反应。Cu缺乏和过量均影响植物正常生长发育, 因此植物进化出精妙复杂的调控网络来严格控制植物体内的Cu含量。植物Cu转运蛋白COPT家族成员与Cu有很高的亲和力, 能够调节植物对Cu的吸收和转运, 在维持植物体内Cu稳态平衡过程中发挥重要作用。COPT蛋白涉及不同的Cu转运功能, 如从外界环境中摄取Cu、从细胞器中输出Cu、长距离运输Cu以及在不同器官间动用和再分配Cu。此外, COPT蛋白在其它离子的稳态平衡维持、昼夜节律性生物钟调控、植物激素合成和植物对激素信号的感受过程中也发挥重要作用。该文综述了模式植物拟南芥(Arabidopsis thaliana) COPT家族各成员的表达和定位、调控机制以及生物学功能等方面的最新进展。

关键词: 铜转运蛋白, COPT, Cu稳态, Cu吸收, 再分配

Abstract: Copper (Cu) is an essential trace element in plants and is involved in many physiological and biochemical reactions as a cofactor of various enzymes. Cu deficiency and excess can affect the normal growth and development, so plants have developed sophisticated regulatory networks to strictly control Cu content. The copper transporter COPT, having high affinity with Cu can regulate the absorption and transport of Cu, and plays an important role in maintaining Cu homeostasis. COPT proteins are involved in different processes of Cu transport, such as uptake of Cu from the external environment, export of Cu from organelles, transport of Cu over long distances, and mobilization and redistribution of Cu between different organs. In addition, COPT proteins play an important role in maintaining homeostasis of other ions, regulating the circadian clock, involving in synthesis of plant hormones and perception of hormone signals. Here we summarize the recent advances in the expression and localization, regulatory mechanisms, and biological functions of COPT proteins in the model plant Arabidopsis thaliana.

Key words: copper transporter, COPT, Cu homeostasis, Cu absorption, redistribution

王惠,李谨谨,徐金玉,刘鹏,张海燕. 拟南芥COPT家族蛋白研究进展. 植物学报, 2019, 54(6): 786-796.

Hui Wang,Jinjin Li,Jinyu Xu,Peng Liu,Haiyan Zhang. Advances in Studies on the COPT Proteins in Arabidopsis thaliana. Chinese Bulletin of Botany, 2019, 54(6): 786-796.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB19045

https://www.chinbullbotany.com/CN/Y2019/V54/I6/786

图/表 2

表1 利用启动子-GUS构建检测的拟南芥COPT家族基因表达模式

Table 1 Expression patterns of COPT family genes detected by use of promoter-GUS in Arabidopsis thaliana

基因名称	表达器官													参考文献
	花				叶				茎		根
	花粉	花丝	雌蕊	胚珠	子叶	表皮毛	保卫细胞	维管组织	维管组织	胚轴	主根	侧根	根毛
COPT1	++	-	/	/	+	+	+	-	-	+	+	+	/	Kampfenkel et al.,1995; Sancenon et al., 2004
COPT2	++	-	-	/	+	+	/	/	-	/	+	+	+	Sancenón et al., 2003; Perea-García et al., 2013
COPT3	++	+	/	/	/	/	/	+	/	/	/	/	/	Andrés-Colás et al., 2018
COPT5	-	+	++	++	+	+	/	++	+	+	++	+	++	Garcia-Molina et al., 2011
COPT6	+	+	+	+	+	+	+	++	++	/	-	+	/	Jung et al., 2012; Garcia-Molina et al., 2013

图1 拟南芥COPT的亚细胞定位与Cu的转运(改自Rodríguez et al., 1999; Balandin and Castresana, 2002; Wintz et al., 2003; Yruela, 2009; Garcia et al., 2014) Cu2+可以被ZIP蛋白吸收或被FRO还原为Cu+, Cu+通过COPT1、COPT2和COPT6蛋白转运进入细胞质。在细胞内, COPT5定位于液泡膜和液泡前体膜, 将液泡和液泡前体中的Cu+转运到细胞质基质中。推测COPT3定位于内质网膜, 并将内质网中的Cu+转运到细胞质基质中。HMA6和HMA1分别定位于叶绿体外膜和内膜, 分别负责将Cu+和Cu2+转运到叶绿体基质。HMA8定位于类囊体膜上, 可将叶绿体基质的Cu+转运到类囊体腔中。HMA7能够转运Cu+通过高尔基体膜, 参与乙烯反应。HMA5定位于细胞膜, 介导Cu+的外排。

Figure 1 Subcellular localization of COPT and copper traffic in Arabidopsis (modified from Rodríguez et al., 1999; Balandin and Castresana, 2002; Wintz et al., 2003; Yruela, 2009; Garcia et al., 2014) Cu2+can be uptaken by ZIP proteins or reduced to Cu+ by FRO and enters into the cytosol through the COPT1, COPT2 and COPT6 transporters. In the cell, COPT5 localizes in the tonoplast and vacuolar precursor membranes and transports Cu+ to the cytosol. It is speculated that COPT3 localizes to the ER membrane and transports Cu+ to the cytosol. HMA6 and HMA1 are localized in the membranes of the chloroplast and responsible for transporting Cu+ and Cu2+ to the cytosol, respectively. HMA8 is located in the thylakoid membrane and transports the Cu+ of the stroma into the thylakoid cavity. HMA7 transports Cu+ through the golgi membrane and participating in the ethylene reaction. HMA5 localizes in the cell membrane and mediates the efflux of Cu+.

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