摘要
镉(Cd)是对植物最具毒害作用的环境污染物之一,它可干扰植物的许多生理生化过程,如干扰光合作用、呼吸作用、氮和蛋白质代谢以及营养元素的吸收。
本研究以10种豆科植物为材料,采用溶液培养法对其进行了Cd耐性的对比研究。种子萌发和幼苗生长的结果表明：绿豆(Phaseolus aureus)是Cd敏感植物,箭舌豌豆(Vicia Sativa)是Cd耐性植物。
研究了10-100μM Cd (CdCl2)(?)(?)种12h对萌发的绿豆和箭舌豌豆种子内以及100μM Cd处理幼苗3d对幼苗叶片中一些生理生化特性的影响。结果表明：≥50μM Cd处理降低了绿豆和箭舌豌豆萌发种子内蛋白酶活性和游离氨基酸含量,但增加了其中的硫代巴比妥酸反应产物(TBARS,Thiobarbituric acid reactive substances)含量；100μMCd降低了二种植物的根系活力、叶片的叶绿素含量及光合作用、硝酸还原酶活性和硝态N、Ca、Mg、Fe、Zn和Mo元素的含量。这些参数的降低在绿豆幼苗中较为明显。
研究了100μM Cd (CdCl2)处理期间绿豆和箭舌豌豆叶的超氧化物歧化酶(Superoxide dismutase, SOD)、抗坏血酸过氧化物酶(Ascorbate peroxidase, APX)、过氧化氢酶(Catalase, CAT)、谷胱甘肽还原酶(Glutathione reductase, GR)和愈创木酚过氧化物酶(Guaiacol peroxidase, GPOD)活性以及还原型谷胱甘肽(Reduced glutathione, GSH)和抗坏血酸(Ascorbic acid, ASC)含量的变化。结果表明：在Cd处理9d期间,绿豆和箭舌豌豆叶的SOD、APX和CAT的活性以及绿豆叶GR活性呈先升后降的趋势,而绿豆和箭舌豌豆叶的GPOD以及箭舌豌豆叶的GR活性呈上升趋势。与此同时,绿豆叶的GSH和ASC含量呈下降的趋势,箭舌豌豆叶的GSH则先稍有下降、后升到对照的水平,ASC含量的变化不明显。与绿豆相比,CAT、APX和SOD活性在箭舌豌豆叶中较高。
采用组织化学染色法观察了100μMCd (CdCl2)处理6d后绿豆和箭舌豌豆幼苗叶内超氧阴离子(O2·-)和过氧化氢(H2O2)的积累。结果发现：Cd处理明显增加了二种植物叶片中O2·-和H202的量、质膜NADPH氧化酶活性以及共质体和质外体中SOD和APX的活性；二种植物叶片中,尤其是绿豆叶片质外体的GPOD明显升高；依赖于H2O2的CeCl3沉淀颗粒主要定位在细胞壁和胞外空间,少量的定位于叶绿体中,依赖于O2·-的二甲基联苯胺(DAB)沉淀颗粒主要定位于叶绿体、细胞壁以及胞外空间,这表明Cd处理使二种植物叶产生了02·-和H2O2,且在绿豆的叶绿体和胞外空间生成的较多。
采用NADPH氧化酶的抑制剂二苯基碘(DPI)和POD的抑制剂叠氮化钠(NaN3)的研究结果表明：在二种植物叶内,Cd诱导积累的ROS主要来源于质膜NADPH氧化酶和质外体过氧化物酶。和箭舌豌豆叶相比,绿豆叶中Cd诱导产生的O2·-和H202、质膜NADPH氧化酶和质外体过氧化物酶活性升高更明显。相比之下,箭舌豌豆叶共质体有较高的SOD和APX活性。
研究了水杨酸(Salicylic acid,SA)对Cd毒害的可能性缓解作用。结果显示：50μM Cd (CdCl2)明显增加了绿豆和箭舌豌豆根系中电解质泄漏率和TBARS含量。100μMSA浸种16h或预处理幼苗16h缓解了Cd对二种植物生长的抑制作用,降低了Cd造成的氧化伤害。SA浸种比SA预处理幼苗对Cd毒害的缓解效果更好。100μM SA浸种降低了O2·-和H2O2在二种植物根细胞壁上的产生,这个作用在绿豆的根中更明显。结果还表明SA的缓解效应可能与SOD和APX活性的上调有关。
Cadmium (Cd) is one of the most toxic environmental pollutants for plants. Cd can interfere with numerous biochemical and physiological processes including photosynthesis, respiration, nitrogen and protein metabolism, and nutrient uptake. In the present study, ten species were compared in solution culture experiments to investigate the resistance of leguminous plants to excessive Cd. Based on the seed germination and seeding growth, common vetch (Vicia Sativa) and mung bean (Phaseolus aureus) were chosen as two different Cd-resistant and Cd-sensitive species.
Compared with control, the seed soaking with10-100μM Cd (CdCl2) for12h significantly decreased protease activities and the contents of free amino acid in the germinating seeds of mung bean and common vetch, and increased the content of soluble protein and the electrolyte leakages of germinating seeds.100μM Cd decreased the root vitality, chlorophyll content, photosynthetic rate, nitrate reductase activity, and concentrations of nitrate nitrogen, Ca, Mg, Fe, Zn and Mo in leaves of both species. The decrease of these parameters was more pronounced in seedlings of mung bean than those in common vetch.
It was studied that the changes in activity of catalase (CAT), ascorbate peroxidase (APX) superoxide, dismutase (SOD), guaiacol peroxidase (GPOD) and glutathione reductase (GR) and contents of reduced glutathione and ascorbic acid in leaves of mung bean and common vetch seedlings under Cd stress. Results showed that the activities of CAT, APX and SOD initially increased and then decreased and the activities of GPOOD significantly increased in leaves of both species during the9-day of100μM Cd treatment. For the activity of GR, similar change was observed in the leaves of mung bean. In the leaves of common vetch, GR activity increased with the increasing Cd exposure time and reduced glutathione initially decreased and then increased. In contrast, the contents of reduced glutathione and ascorbic acid decreased with Cd treatment time in mung bean. Compared with those in mung bean, the activities of CAT, APX and SOD were higher in common vetch leaves than in mung bean leaves under Cd stress.
Excess Cd treatment induced oxidative stress. The effects of Cd on the accumulation of H2O2and O2'-in leaves of mung bean and common vetch were investigated using both histochemical and cytochemical methods. Cadmium at100μM obviously increased the production of O2·-and H2O2, and the activities of plasma membrane-bound NADPH oxidases and the symplastic and apoplastic activities of SOD and APX in the leaves of both species. Apoplastic GPOD activity was significantly induced in the leaves of both species, particularly in mung bean with100μM Cd. In the leaves of both species treated with100μM Cd, H2O2-dependent CeCl3precipitate was mainly localized in cell walls and the corners of the extracellular space as well as a small amount in chloroplasts of the mesophyll cells, and O2·-dependent DAB precipitates were observed mainly in chloroplasts, cell walls, and extracellular spaces. It was evident that more H2O2and O2·-were produced in the chloroplasts and extracellular spaces of mung bean than in those of common vetch. Experiments with diphenyleneiodonium as an inhibitor of NADPH oxidase and NaN3as an inhibitor of peroxidase showed that the majority of Cd-induced ROS production in the leaves of both species may be from plasma membrane-bound NADPH oxidase and apoplastic GPOD. Compared with those in the species common vetch, Cd-induced O2·-and H2O2production, NADPH oxidase and apoplastic GPOD activity increase were more pronounced in mung bean. In contrast, common vetch had higher leaf symplastic SOD and APX activities than mung bean.
The possible mediatory role of salicylic acid (SA) in protecting plants from Cd toxicity was studied. Treatment with50μM Cd significantly increased the electrolyte leakages and total thiobarbituric acid reactive substances content in roots of mung bean and common vetch. Seeds soaking with100μM SA for16h or seedlings pretreatment with100μM SA for16h alleviated the negative effect of Cd on growth of both species. The alleviation of Cd toxicity was more pronounced by seed soaking with SA than seedlings pretreatment with SA. Seeds soaking with100μM SA decreased the production of O2·-and H2O2in root apoplast of both species. This effect was more evident in the roots of mung bean than in roots of common vetch. The data suggest that the beneficial effect of SA during an earlier growth could be related to up-regulation of antioxidant enzyme SOD and APX.
引文
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