Research Progress on the Interaction Mechanism between Hyperaccumulator and Heavy Metals and Its Application
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摘要: 社会发展过程中对矿产资源的勘查和开采利用所带来的重金属污染已对生态系统和人类健康造成严重威胁。超富集植物对重金属具有超富集、超耐受能力,是降低环境重金属污染、保障人类健康、实现绿色矿产勘查的有效途径,在植物修复、植物采矿和植物找矿中已获得了广泛应用。深入探索超富集植物的富集和耐受机制,揭示重金属-植物相互作用规律,提高植物对重金属的富集能力,是当前国际上研究热点。本文在简要介绍重金属对植物作用的基础上,阐述了重金属诱导氧化应激机制,重点关注重金属超富集植物富集机理研究,对其在解毒和耐受机制等领域的研究进展进行了评述。当前研究认为:①对超富集植物而言,根系分泌物与根际微生物的共同作用促进了重金属溶解,经共质体、质外体途径吸收后,重金属通过木质部向上转运,并隔离在液泡中,实现对重金属的超富集;②重金属通过与小分子有机酸、细胞壁、植物螯合肽结合,以及液泡隔离,可降低细胞质中游离金属离子浓度,增强植物耐受性;③重金属胁迫下,植物将激活多种特异性抗氧化酶,抵御氧化应激反应,实现对重金属的超耐受。④本文分析认为,植物中砷诱导的氧化应激反应机制可能是由砷的还原与甲基化过程及Haber-Weiss反应三部分构成。对重金属超富集植物的富集与耐受过程所涉及的生理与生化作用进行深入研究,揭示关键性影响因素与相关规律,寻找提升其特异性富集与指示能力的有效途径,将有助于超富集植物研究与应用向纵深发展。Abstract:
BACKGROUND Heavy metal pollution caused by the exploitation of mineral resources in the social development has caused serious threats to the ecosystem and human health. Hyperaccumulating plants have super-enrichment and super-tolerance capabilities for heavy metals, which is an effective way to reduce environmental heavy metal pollution, protect human health, and realize green mineral exploration. It has been widely used in phytoremediation, plant mining and plant prospecting. OBJECTIVES To better understand the enrichment and tolerance mechanisms of hyperaccumulating plants, reveal the principles of heavy metal-plant interactions, and improve the ability of plants to accumulate heavy metals. METHODS Based on a brief description of the effects of heavy metals on plants, the focus of this article is the accumulation mechanism of heavy metal hyperaccumulation plants, and a review of the progress in the fields of detoxification and tolerance mechanisms. RESULTS (1) The root exudates of hyperaccumulator and microorganisms work together to promote the dissolution of heavy metals. After being absorbed by the symplastic and apoplastic pathways, the heavy metals are transported upwards to aerial parts through xylem, and segregated in vacuoles, achieving the hyperaccumulation of heavy metals. (2) Concentration of free metal ions in cytoplasm can be reduced by combining heavy metals with small molecular organic acids, cell walls, phytochelatins and vacuole isolation, which increases plant tolerance. (3) Under heavy metal stress, plants activate a variety of specific antioxidant enzymes to resist oxidative stress and achieve hypertolerance on heavy metals. (4) A possible mechanism is suggested that arsenic-induced oxidative stress in plants should be composed of arsenic reduction and methylation, and Haber-Weiss reaction. CONCLUSIONS In-depth research on the physiological and biochemical processes involved in hyperaccumulation and hypertolerance of hyperaccumulator reveals key factors and related principles. Finding effective ways to improve their specific accumulation and indication capabilities will contribute to the research and application of hyperaccumulators to develop in depth. -
图 2 植物体内重金属元素超富集的主要过程与机制[50]
Figure 2.
图 3 重金属液泡积累机制,液泡膜转运蛋白直接将重金属元素带入液泡[99]
Figure 3.
表 1 植物中微量元素的作用与机制
Table 1. Function and mechanism of micronutrients in plants
微量元素 作用(适量) 过量
(干重浓度)毒性机制 毒性症状 Zn[13] 促进植物形成花粉;氧化还原酶、转移酶、
水解酶、裂解酶、异构酶、连接酶组成成分>20mg/kg 增强脂质过氧化酶活性 引起遗传变异,抑制植物
生长Cu[13] 多种酶辅因子;参与细胞壁代谢,叶绿体、
线粒体电子传递;氧化磷酸化和铁动员,
氮同化,脱落酸合成等>20μg/g 抑制酶活性及蛋白质
功能,引起氧化应激抑制胚芽发育、种子活性、
植株发育,导致萎黄、坏死Mn[13] 参与ATP酰基脂质、蛋白质、脂肪酸生物
合成,参与RuBP羧化酶反应,光合作用
的氧化还原过程>(10~100)μg/g 干扰其他营养元素吸收
利用,诱导氧化应激影响能量代谢,降低光合
作用速率Se[24, 28] 保护细胞膜,防止不饱和脂肪酸氧化 >2mg/kg 非特异性硒蛋白积累,
氧化应激抑制植物生长 Ni[29-30] 脲酶组成成分 >(10~50)mg/kg 破坏叶绿素结构,降低
叶绿素含量抑制光合作用、氮代谢、
酶活性,产生活性氧,生长
速率下降Co[30] 诱导淀粉积累 >368mg/kg 破坏叶绿素结构,降低
叶绿素含量降低生长速率,光合作用
速率下降表 2 在不同培养条件下生长的不同植物中,重金属激活抗氧化酶差异
Table 2. Heavy metal-induced activation of antioxidant enzymes in different plant species grown in different condition
植物名称 胁迫元素与浓度 胁迫时间 基质 抗氧化酶 活性变化情况 江南星厥[109]
(Microsorum fortunei)Cd:1000μmol/L 15d 水培 CAT、POD、GST、CCP 降低 水合欢[110]
(Neptunia olerace)Cd:50、100、180mg/kg Pb:500、1000、1800mg/kg 37d 土培 根中CAT、茎中SOD、叶中POD 先降低后升高 根和茎中SOD、茎中CAT、叶中POD 大聚藻[111]
(Myriophyllum aquaticum)Cd:10、20、40、80、160mg/kg 28d 水培 SOD、POD、PRO 升高 印度芥菜[112] Cu、Zn、Pb、Cd:50μmol/L 96h 水培 SOD、CAT、APX 根部升高,地上部分先升高后降低(高于对照组) 墨旱莲[113]
(Eclipta prostrata)Pb:100、200、400、800、1600mg/kg 30d 土培 SOD、CAT、APX、GR 升高 印度芥菜、紫花苜蓿[114] Cd:7 5、150、300、600mg/kg 14d 土培 SOD 紫花苜蓿茎中升高 CAT 两者根、茎中均有升高 注:CAT—过氧化氢酶(catalase);POD—过氧化物酶(peroxidase);GST—谷胱甘肽硫转移酶(glutathione-S-transferase);CCP—细胞色素c过氧化物酶(cytochrome c peroxidase);SOD—超氧化物歧化酶(superoxide dismutase);PRO—脯氨酸(proline);APX—抗坏血酸过氧化物酶(ascorbate peroxidase);GR—谷胱甘肽还原酶(glutathione reductase)。 表 3 部分重金属超富集植物及其富集能力和应用
Table 3. Accumulation ability of several hyperaccumulators and their application
元素 超富集植物名称 生长条件 植物中重金属浓度(干重) 应用 Ni 褐蓝菜[75] 100μmol/L水培,>60天 根:33.6μmol/g
叶:130μmol/g/ Odontarrhena chalcidica
(庭芥属)[116]蛇纹石(serpentine)土壤,6.5个月 地上部分:13μg/g 植物采矿 小野芥菜[116]
(Noccaea goesingensis)蛇纹石土壤,11个月 地上部分:8μg/g 植物采矿 Senecio conrathii(菊科)[117] 总Ni:503μg/g,可溶性
Ni:0.1μg/g,土壤叶:1558μg/g 植物修复 髭脉桤叶树[118]
(Clethra barbinervis)500μmol/L水培,12周 根:1310μg/g
茎:542μg/g
叶:804μg/g/ 少根紫萍[30]
(Landoltia punctata)10mg/L水培,10天 叶:2013mg/kg 植物修复 Co 髭脉桤叶树[118] 500μmol/L水培,12周 根:1810μg/g
茎:246μg/g
叶:1770μg/g/ 少根紫萍[30] 10mg/L水培,10天 叶:1998mg/kg 植物修复 Cd 伴矿景天[82] 总Cd:36~157mg/kg,
CaCl2可提取态Cd:
1.83~14.2mg/kg,土壤地上部分:574~1470mg/kg 植物修复 多穗稗[65] 100mg/L水培,62天 根:299mg/kg
叶:233mg/kg植物修复 东南景天[79] 25μmol/L水培,30天 根:150μg/g;
地上部分:500μg/g植物修复 宝山堇菜[119]
(Viola baoshanensis)100μmol/L水培,2天 根:3500mg/kg
地上部分:1750mg/kg植物修复 狐尾藻[111]
(Myriophyllum aquaticum)40mg/L水培,28天 17970mg/kg 植物修复 龙葵[66] 20mg/kg,土壤,14天 根:95μg/g
地上部分:128μg/g植物修复 Zn 褐蓝菜[75] 100μmol/L水培,60天 根:46.4μmol/g
叶:161μmol/g(干重)/ 伴矿景天[82] 总Zn:1930~6200mg/kg,
CaCl2可提取态
Zn:24~162mg/kg,土壤地上部分:9020~14600mg/kg 植物修复 As 凤尾蕨[120] 10mg/kg,土壤,30天 根:1885mg/kg
叶:2562mg/kg植物修复 Pb 墨旱莲[113]
(Eclipta prostrata)1600mg/kg,土壤,30天 根:7229μg/g
地上部分:12484μg/g植物修复 -
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