Research Advances on Plant Root Exudates in Response to Heavy Metal Stress
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摘要: 植物在遭受重金属胁迫时，会改变根系分泌物的分泌情况来响应重金属胁迫。根系分泌物的变化与植物种类、重金属类型等多种因素有关。本文综述了重金属胁迫下，植物根系分泌物中有机酸、氨基酸、可溶性糖的变化以及这些组分的作用，为研究植物对重金属的耐受性和适应性，以及明确各组分的响应机制，提供理论依据。Abstract: In response to heavy metal stress plants alter the root exudate secretion in ways depending on the plant species and type of heavy metals, among other factors. This article reviews the published literature on the organic acids, amino acids, and soluble sugars in root exudates and their roles in the chemistry and physiology of the plants under varieties of heavy metal stresses. The referred material and documents concerning the tolerance and adaptability of plants to heavy metals as well as the response mechanisms involving the key chemicals would provide a concise and update information for the scientists interested in the field of study.
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Key words:
- Soil /
- heavy metal /
- stress /
- root exudates
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图 1 植物根系分泌物活化土壤重金属的途径
Figure 1. Activating pathways of heavy metals in soil by plant root exudates
图 2 部分氨基酸以及与重金属相关的氮代谢物合成途径
Figure 2. Synthesis pathways of some amino acids and nitrogen metabolites related to heavy metals
表 1 根系分泌物的分类依据
Table 1. Basis for classification of root exudates
分类方式
Classification method种类
Type特点
Characteristic
按来源分类
Classification by source分泌物
Secretion根系细胞主动释放的化学物质
Chemicals actively released by root cells渗出物
Exudate根系细胞通过浓度差扩散作用释放出的小分子化合物
Small molecular compounds released by root cells through concentration difference diffusion黏胶质
Viscose substances根冠细胞、未形成次生壁的表皮细胞、根毛分泌的粘胶状物质
Root cap cells, epidermal cells without secondary wall and mucilaginous substances secreted by root hairs裂解物
Lysate从破裂的或者已经死亡细胞中释放出来的化学物质
Chemicals released from ruptured or dead cells
按性质分类
Classification by nature脱落物及溶解产物
Shedding and dissolved products植物根系在生长过程中脱落的根冠、根表皮、根毛细胞等
The root cap, root epidermis and root hair cells shed by plant roots during growth高分子凝胶状物质
Polymeric gelatinous material黏胶物质、黏液、边缘细胞、多聚糖和糖醛酸、各种酶类
Viscose substances, mucus, marginal cells, polysaccharides and uronic acids, various enzymes小分子有机化合物
Small molecule organic compound有机酸、糖类、氨基酸、酚类、多肽、部分无机离子等
Organic acids, sugars, amino acids, phenols, polypeptides, some inorganic ions, etc/
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表 2 植物受重金属胁迫时常见的根系分泌物
Table 2. Common root exudates of plants under heavy metal stress
类别
Category常见的组分
Common components有机酸
Organic acid草酸、柠檬酸、酒石酸 、乳酸、乙酸、琥珀酸、苹果酸、丙二酸、乙醇酸、丙酮酸等
Oxalic acid, citric acid, tartaric acid, lactic acid, acetic acid, succinic acid, malic acid, malonic acid, glycolic acid, pyruvic acid, etc.氨基酸
Amino acid丙氨酸、缬氨酸、甘氨酸 、谷氨酸、苏氨酸、γ-氨基丁酸、脯氨酸 、蛋氨酸、谷氨酰胺、组氨酸、天冬氨酸、精氨酸、丝氨酸、天冬酰胺、半胱氨酸、苯丙氨酸等
Alanine, valine, glycine, glutamic acid, threonine, γ- aminobutyric acid, proline, methionine, glutamine, histidine, aspartic acid, arginine, serine, asparagine, cysteine, phenylalanine, etc.可溶性糖
Soluble sugar果糖、葡萄糖、蔗糖、半乳糖、木糖、阿拉伯糖、乳糖、鼠李糖、岩藻糖等
Fructose, glucose, sucrose, galactose, xylose, arabinose, lactose, rhamnose, fucose, etc.
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表 3 不同金属胁迫下植物根系分泌的有机酸类别
Table 3. Types of organic acids secreted by plant roots under heavy metal stresses
植物
Plant金属
Metal有机酸
Organic acid辣椒
PepperCd 草酸、酒石酸和乙酸[65]
Oxalic acid, tartaric acid and acetic acid食用苋菜
Three-colored amaranthCd 琥珀酸、乙酸、苹果酸、草酸和柠檬酸[66]
Succinic acid, acetic acid, malic acid, oxalic acid and citric acid小麦
WheatCd 草酸[67]
Oxalic acid玉米、高粱
Corn and sorghumCd 苹果酸、柠檬酸[18]
Malic acid, citric acid蓖麻
CastorCu 柠檬酸、酒石酸、草酸[62]
Citric acid, tart
aric acid, oxalic acid稗草
BarnyardgrassCd、Cu、Pb复合胁迫
Cd, Cu, Pb compound stress柠檬酸[68]
Citric acid早熟禾、苜蓿、
锦葵
Bluegrass, alfalfa 、mallowCu、Cr、Zn复合胁迫
Cu, Cr, Zn compound stress草酸、苹果酸、富马酸[49]
Oxalic acid, malic acid and fumaric acid菠菜
SpinachAl 草酸[69]
Oxalic acid黑麦
RyeAl 苹果酸[70]
Malic acid大豆
SoybeanAl 苹果酸、草酸[71]
Malic acid, oxalic acid常绿杨
Evergreen poplarAl 草酸、柠檬酸[32]
Oxalic acid, citric acid苔草
CarexPb 草酸、苹果酸、柠檬酸[25]
Oxalic acid, malic acid, citric acid金丝草
Golden silk grassPb 草酸、柠檬酸和苹果酸[21]
Oxalic acid, citric acid and malic acid茶树
Tea plantPb 苹果酸、琥珀酸[23]
Malic acid, succinic acid
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表 4 部分氨基酸在植物受重金属胁迫时的作用
Table 4. Effects of some amino acids on plants under heavy metal stress
氨基酸
Amino acid主要作用
Effectγ-氨基丁酸和丙氨酸
γ-aminobutyric acid and alanine调节维持细胞的pH[36, 77]
Regulate and maintain cell pH甘氨酸
Glycine参与植物螯合肽和抗氧化酶的合成[77]
Involved in the synthesis of plant chelating peptides and antioxidant enzymes甘氨酸和谷氨酸
Glycine and glutamate促进植物体内硝酸盐的还原[78]
Promote the reduction of nitrate in plants天冬氨酸
Aspartic acid与重金属离子结合形成金属复合物[78]
Combine with heavy metal ions to form metal complexes脯氨酸
Proline具有抗氧化的作用[79]
Plays an antioxidant role半胱氨酸、亮氨酸和蛋氨酸
Cysteine, leucine and methionine在植物细胞壁适应胁迫中起着重要作用[80]
Plays an important role in plant cell wall adaptation to stress亮氨酸、异亮氨酸和缬氨酸
Leucine, isoleucine and valine维持细胞渗透平衡[81-82]
Maintain cell osmotic balance
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