生物技术通报 ›› 2022, Vol. 38 ›› Issue (6): 13-26.doi: 10.13560/j.cnki.biotech.bull.1985.2021-1286
刘自然1(), 甄珍1, 陈强2, 李玥莹1, 王泽1, 逄洪波1()
收稿日期:
2021-10-09
出版日期:
2022-06-26
发布日期:
2022-07-11
作者简介:
刘自然,女,硕士研究生,研究方向:作物资源开发与利用;E-mail: 基金资助:
LIU Zi-ran1(), ZHEN Zhen1, CHEN Qiang2, LI Yue-ying1, WANG Ze1, PANG Hong-bo1()
Received:
2021-10-09
Published:
2022-06-26
Online:
2022-07-11
摘要:
镉(Cd)是一种毒性极强的重金属污染物,位于土壤无机污染物首位。Cd因具有强流动性和溶解性而容易被植物体吸收,不仅影响植物正常生理代谢并降低作物品质,还能通过食物链的传递和富集危害人体健康,因此植物对Cd胁迫的响应机制已成为研究的热点。本文综述了Cd胁迫对植物生理代谢和分子层次危害的同时,总结了植物在Cd胁迫下进化出的耐受性机制,以期为培育低积累作物和超积累Cd污染土壤修复植物提供理论依据。
刘自然, 甄珍, 陈强, 李玥莹, 王泽, 逄洪波. 植物响应Cd胁迫研究进展[J]. 生物技术通报, 2022, 38(6): 13-26.
LIU Zi-ran, ZHEN Zhen, CHEN Qiang, LI Yue-ying, WANG Ze, PANG Hong-bo. Research Progress in Plant Response to Cd Stress[J]. Biotechnology Bulletin, 2022, 38(6): 13-26.
图1 植物响应Cd胁迫的生理及分子机制 A:根际分泌物(主要包括高分子的植物螯合剂和金属硫蛋白以及低分子的H+、HCO3-、苹果酸和柠檬酸等)可以结合土壤中游离Cd并酸化根际土壤,是植物阻止Cd进入原生质体的第一道防线;B:植物阻止Cd进入原生质体的第二道防线是利用细胞壁结构物质纤维素、果胶和木质素固定Cd;C:由于植物细胞不具有Cd特异性转运蛋白,所以Cd2+必须与其他必需二价金属离子(如Fe2+/Ca2+/Mn2+)竞争转运蛋白(ZNT1、IRT1/2、NRAMP1/4及HMA4)从而进入原生质体;Cd进入细胞后可能与PCs(植物螯合肽)、MTs(金属硫蛋白)和NA(烟草胺)等螯合剂结合,转运蛋白可以将Cd及其螯合物转运至液泡(MTP1、YSL1、ABCC、YCF1、CAX1/2/4及HMA1/2/3)、叶绿体(ATM3及YSL6)及高尔基体(MTP11)中暂时储存,或排出细胞(PDF8)由木质部利用根压和蒸腾运输到地上部进行储存;D:抗氧化系统(酶类和非酶类系统)负责清除Cd毒害引发的大量活性氧,防止其扰乱细胞内部稳态平衡。酶类系统包括SOD(超氧化物歧化酶)、CAT(过氧化氢酶)、POD(过氧化物酶,peroxidase)、APX(抗坏血酸过氧化物酶)和GR(谷胱甘肽还原酶),非酶类系统包括低质量的代谢物,如GSH(谷胱甘肽)、AsA(抗坏血酸)和脯氨酸等
Fig.1 Physiological and molecular mechanisms of plant response to Cd stress A:Rhizosphere exudates are the first line of defense for plants to prevent Cd from entering the protoplasm. It mainly includes high molecular weight plant chelating agents and metallothionein,as well as low molecular weight H+,HCO3-,malic acid and citric acid,etc. B:The second defense line for plants to prevent Cd from entering protoplasts is to fix Cd with cell wall structural materials cellulose,pectin and lignin. C:Cd2+ must compete with other essential divalent metal ions,such as Fe2+/Ca2+/Mn2+ for transporters(ZNT1,IRT1/2,NRAMP1/4 and HMA4)to enter the protoplast because there is no Cd-specific transporter in plant cells;Cd may bind to chelating agents such as PCs(phytochelatins),MTs(metallothioneins)and NA(nicotinamide)after it enters the cell. Transport proteins can transport Cd and its chelates to the vacuole by MTP1,YSL1,ABCC,YCF1,CAX1/2/4 and HMA1/2/3,to chloroplasts by ATM3 and YSL6,and to Golgi by MTP11 temporarily stored,or Cd is excreted from cells by PDF8 and is transported from the xylem to the ground for storage using root pressure and transpiration. D:Antioxidant systems(such as enzymatic and non-enzymatic oxidants)are crucial for the detoxification of the ROS under stress conditions. It is responsible for preventing it from disturbing the homeostasis of cells. Enzymatic systems in plants include SOD(superoxide dismutase),CAT(catalase from micrococcus lysodeiktic),POD(peroxidase),APX(ascorbate peroxidase)and GR(glutathione reductase). The non-enzymatic antioxidant system includes low-mass metabolites like GSH(glutathione glutathione),AsA(ascorbic acid),and proline
图2 7种常见的重金属转运蛋白结构图 直线:质膜和液泡膜等膜系统;长方形:跨膜结构域;曲线:跨膜区间。A:Zn/Fe转运蛋白(ZIP);B:天然抗性相关巨噬细胞蛋白(NRAMP);C:重金属ATP酶(HMA);D:金属耐受蛋白(MTP);E:阳离子交换体(CAX);F:ATP结合盒转运蛋白(ABCC),F1:ABCC半转运子,F2:ABCC全转运子;G:黄色条纹转运蛋白(YSL)
Fig.2 Structures of 7 common heavy metal transporters Straight line:Membrane systems such as plasma membrane and vacuolar membrane. Rectangle:Transmembrane domain. Curve:Transmembrane interval. A:Zinc and iron transporter proteins(ZIP). B:Natural resistance-associated macrophage proteins(NRAMP). C:Heavy metal ATPase(HMA). D:Metal tolerance protein(MTP). E:Cation exchanger(CAX). F:ATP-binding cassette transporter(ABCC);F1:ABCC semit ransporter;F2:ABCC full transporter. G:Yellow stripe-like transporter(YSL)
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