稀脉萍（Lemna aequinoctialis）和紫萍（Spirodela polyrrhiza）的重金属生态毒理学研究
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摘要
生态毒理学是上世纪80年代前后才形成的新学科，许多方面尚处于完善过程中，而在我国，生态毒理学研究还属于刚起步，相对于其它研究显得尤为薄弱。加强这方面的研究不仅可以促使我国在该学科的上的进步，而且在促进我国经济发展模式向可持续发展转变上有积极的推动作用。
生态毒理学是建立在生态学基础的毒理学，着重研究研究外源化学物质，尤其是有毒化学品对生态系统结构和功能的危害作用及生态系统所发生的反应，也探索这些物质在生态系统中的迁移转化规律。同时，还为有毒物质污染环境的生态(生物)恢复、生态风险评价提供技术支持和理论指导。其中，重金属作为一类特殊的有毒物质受到生态毒理学研究者的特别关注。生态毒理学给毒理学研究带来了深刻的变革，也改变了许多环境技术的发展方向。
重金属对植物的生态效应是多方面的，既有个体水平上的，也有更高水平上的，如种群水平和群落水平上的，既有生理生化方面的，也有遗传演化方面的，但植物也能够产生多种耐性机制以减轻重金属的毒害效应，如产生胁迫蛋白、胁迫乙烯，启动抗氧化保护系统，合成金属硫蛋白、植物螯合素，产生应激乙烯，增加排除量，将重金属集中于局部部位并隔离起来等。理解重金属的毒性及其产生机制，了解植物对胁迫的反应，既可以增加我们对重金属的生态效应的认识，还能够帮助我们更深入地了解重金属的不良的生态学效应，促使我们更加合理地使用重金属，以尽量减轻重金属对生态系统的不利影响。
各种化学品，特别是像重金属及其制品、农药、除草剂之类的有毒化学物质对生态系统的结构和功能有很大的危害性，对这种危害性进行评价是进行有毒化学品科学管理的基础，也是保护环境，维护生态安全的有效手段。发达国家不仅有成熟的标准评价方法，而且有相应的法规保证。我国在这方面还比较落后，但严峻的环境形势迫使我们必须加快研究步伐，尤其是应该将国外成熟的评价方法能尽快本土化，以便为加强有毒化学品管理的立法、执法和科学管理提供科学依据。国外在使用维管束植物进行毒性评价时，浮萍属的植物是其中研究和应用最多的，加快本土化的步伐会对保护我国的环境产生积极影响。
本论文利用我国浮萍科植物中分布最广的两个种类作为测试材料，从几个方面对自然
来源的稀脉萍(L。。na aoquj，] ocrj;/js)和紫萍(sPjrod。/刀夕01少r厂hiz召)的重金属生
态毒理学进行了较全面的研究，其目的是建立以本土浮萍种类为标准植物毒性m.J试材料的
国家标准方法，发挥浮萍在水体污染植物修复中的作用.在研究中有新的发现，并探讨了
其机理.研究结果不仅增加了人们对重金属毒性效应的认识，也可以为建立我国利用本土
浮萍属植物进行有毒有害物质的植物毒性评价的标准方法，充分发挥其在重金属污染的生
态恢复中的作用提供科学依据.主要工作如下:
1.在实验中研究7重金属锡(Cd)、铜(Cu)、锌(Zn)、镍(Ni)、汞(Hg)、铬(Cr)胁
迫对自然来源的稀脉萍(L.刁四u，’刀octf召1j’s)和锡(cd)胁迫对自然来源的紫萍(s.
Po了厂rh，’z刀)的效应，发现它们能够产生使这两种浮萍的子体与母体提前分离，致使
群体发生解体的效应，也发现锡和铜的这种效应非常灵敏，并呈现明显的时间和浓度
依赖关系，其它四种金属的这种效应要低于铜和锡，时间和浓度依赖关系也比较弱.
并且，对稀脉萍，这种效应在最快时暴露于锡或铜处理下2一3个小时就可达到显著水
平，2礴小时即可完成，而对紫萍需要30小时才脆达到显著水平，72小时才基本完成。
同时还证明铅(Pb)对自然来源的稀脉萍(乙.五时ul’肋cti左l，’，.)不产生这种效应。分
析了过去的研究没有报道这种现象的原因可能是因为他们使用的材料多是经过特殊筛
选的浮萍的特殊品系，遗传多样性水平大大降低，已经丢失了产生这种反应的敏感株
系，或者是他们忽略了这种反应，也可能是由浮萍的地理差异导致的遗传上的原因.
认为能够形成浓度依赖和时间依赖关系也是由于自然来源的稀脉萍的遗传多样性较高
的缘故，使得敏感个体和杭性个体构成正态分布，是形成时间一浓度一效应关系的基础.
并推测两种浮萍对重金属的这种反应对植物适应环境变化有一定价值，也可以作为重
金属植物毒性评价的生物标记。重金属锅过量会对植物的生存形成多种胁迫效应，如
生长率的下降，光合作用效率降低，呼吸强度增加等，但它们能够促使稀脉萍和紫萍
群体发生解体的现象尚未见文献报道。
2.以铜、锡的胁迫效应为例，进一步研究了重金属产生的这种效应的机制，并测定了锅
胁迫下紫萍的一些生理生化指标的变化，较全面地了解了锅对紫萍的毒性效应。气相
色谱分析表明，在胁迫培养过程中，乙烯的产生量有很大增加，远远超过正常生长时
的产生量.说明在它们的胁迫下，有胁迫乙烯产生.根据已有的对其它植物进行研究
II
的报道，锅、铜胁迫能够刺激这些植物产生胁迫乙烯，并结合乙烯在植物体内具有的
能够加快衰老、促进脱叶的生理学作用，推侧?
Ecotoxicology is a newsubjecl just founded in about 1980 and is developing rapidlyOver the past two decades, ecotoxicology has experienced a rapid progress,thus it has become to be very important in protecting ecological safety from toxic contaminants. In our country, however, ecotoxicology is in its babyhood, to which we do not know very clear in many aspects. Enhancing the research in this field not only can faditate the advancement of this new subject in our country, but also greatly hasten the conversion of the economic pattern of our country into sustainable development.
Based on ecology, ecotoxicology is to study the hazardous influences of contaminants, especially, toxic chemicals on the structure and functions of ecosystem, the response of ecosystems to them, and the pathways of the toxicants moving, transforming and decomposing in the surrounding. Also, acotoxicology provides the technological base and tboretic steering for the ecological remediation of contaminated environment and ecological risk assessment. Of the toxic chemicals, heavy metals are given special attention by ecotocicological researchers because they are important inorganic poIlutantsEcotoxicology brought a profound revolution in toxicology, and led a change of the direction of the advance ofome environmentaltechniques.
Heavy metals have lots of effects on the plants, not only on the individuals, and on the populations and the comnunities in higher levels, but also in physiological and biochemical aspects and in genetics and evolution, e.g. in physiology, depressing the activity of some enzymes, initiating a few free radicals, reducing the growth of plants, degrading chlorophyll, inhibiting the elongation of the root and so on. But, the plant also develops plenty of mechanisms to resist the toxicities of heavy metals, such as stimulating stress proteins and stress ethylene production, activating antioxidant system to eliminate the fe radicals, synthesizing metallothioneins-like and phytochelatins, excluding more metals and accumulating and compartmentalizing in some organs. To understand the toxicity and mechanisms underlying of the heavy metals can increase our knowledge about thai ecological effects, and facilitate us to know their hazardous ecological effects more clear, which would urge us to use them
more availably and rationally so that their negative effects could be alleviated as possible.
Various chemicals, especially toxc chemicals, such as heavy metals or heavy metals-contained, insecticide, herbicide and so on, have great toxic effects orthe structure and functions ofinosphere. To assess their toxicity, including phytotoxicity, is the basis of scientific management fa them, and is profound to protect environmental quality and ecological safety.
Developed countries not only have weHestablished standard method to process the toxicity evaluation of toxicants, but also many related laws and rules for it. Our country drps greatly behind them in this aspect. However, the environmental status of our country has been so severe that we have to enhance the research in this field. It should be a short way to study the feasibility of using indigene organisms in assessing the taicity of pollutants based on using the established standard method. Management for reference, legislation and executing the law of toxic chemicals could be on the scientific basis when standard toxicity assessment method of our country is established. By far, the most commonly used vascular plants in toxicity tests are duckweeds, members of Lemanaceae.
This thesis conducted a completely ecotoxicological studyon the effects of heavy metals on two duckweed species, most common in our country,recollected from natural populations instead of from special strains selected in laboratory with axentic conditionsThe objects are to provide an available autochthonous model testing material for phytotoxicity assessment ofiiic chemicals in our country, an excellent plant material for phytoremediation of cadmimpolluted water. In
生态毒理学是建立在生态学基础的毒理学，着重研究研究外源化学物质，尤其是有毒化学品对生态系统结构和功能的危害作用及生态系统所发生的反应，也探索这些物质在生态系统中的迁移转化规律。同时，还为有毒物质污染环境的生态(生物)恢复、生态风险评价提供技术支持和理论指导。其中，重金属作为一类特殊的有毒物质受到生态毒理学研究者的特别关注。生态毒理学给毒理学研究带来了深刻的变革，也改变了许多环境技术的发展方向。
重金属对植物的生态效应是多方面的，既有个体水平上的，也有更高水平上的，如种群水平和群落水平上的，既有生理生化方面的，也有遗传演化方面的，但植物也能够产生多种耐性机制以减轻重金属的毒害效应，如产生胁迫蛋白、胁迫乙烯，启动抗氧化保护系统，合成金属硫蛋白、植物螯合素，产生应激乙烯，增加排除量，将重金属集中于局部部位并隔离起来等。理解重金属的毒性及其产生机制，了解植物对胁迫的反应，既可以增加我们对重金属的生态效应的认识，还能够帮助我们更深入地了解重金属的不良的生态学效应，促使我们更加合理地使用重金属，以尽量减轻重金属对生态系统的不利影响。
各种化学品，特别是像重金属及其制品、农药、除草剂之类的有毒化学物质对生态系统的结构和功能有很大的危害性，对这种危害性进行评价是进行有毒化学品科学管理的基础，也是保护环境，维护生态安全的有效手段。发达国家不仅有成熟的标准评价方法，而且有相应的法规保证。我国在这方面还比较落后，但严峻的环境形势迫使我们必须加快研究步伐，尤其是应该将国外成熟的评价方法能尽快本土化，以便为加强有毒化学品管理的立法、执法和科学管理提供科学依据。国外在使用维管束植物进行毒性评价时，浮萍属的植物是其中研究和应用最多的，加快本土化的步伐会对保护我国的环境产生积极影响。
本论文利用我国浮萍科植物中分布最广的两个种类作为测试材料，从几个方面对自然
来源的稀脉萍(L。。na aoquj，] ocrj;/js)和紫萍(sPjrod。/刀夕01少r厂hiz召)的重金属生
态毒理学进行了较全面的研究，其目的是建立以本土浮萍种类为标准植物毒性m.J试材料的
国家标准方法，发挥浮萍在水体污染植物修复中的作用.在研究中有新的发现，并探讨了
其机理.研究结果不仅增加了人们对重金属毒性效应的认识，也可以为建立我国利用本土
浮萍属植物进行有毒有害物质的植物毒性评价的标准方法，充分发挥其在重金属污染的生
态恢复中的作用提供科学依据.主要工作如下:
1.在实验中研究7重金属锡(Cd)、铜(Cu)、锌(Zn)、镍(Ni)、汞(Hg)、铬(Cr)胁
迫对自然来源的稀脉萍(L.刁四u，’刀octf召1j’s)和锡(cd)胁迫对自然来源的紫萍(s.
Po了厂rh，’z刀)的效应，发现它们能够产生使这两种浮萍的子体与母体提前分离，致使
群体发生解体的效应，也发现锡和铜的这种效应非常灵敏，并呈现明显的时间和浓度
依赖关系，其它四种金属的这种效应要低于铜和锡，时间和浓度依赖关系也比较弱.
并且，对稀脉萍，这种效应在最快时暴露于锡或铜处理下2一3个小时就可达到显著水
平，2礴小时即可完成，而对紫萍需要30小时才脆达到显著水平，72小时才基本完成。
同时还证明铅(Pb)对自然来源的稀脉萍(乙.五时ul’肋cti左l，’，.)不产生这种效应。分
析了过去的研究没有报道这种现象的原因可能是因为他们使用的材料多是经过特殊筛
选的浮萍的特殊品系，遗传多样性水平大大降低，已经丢失了产生这种反应的敏感株
系，或者是他们忽略了这种反应，也可能是由浮萍的地理差异导致的遗传上的原因.
认为能够形成浓度依赖和时间依赖关系也是由于自然来源的稀脉萍的遗传多样性较高
的缘故，使得敏感个体和杭性个体构成正态分布，是形成时间一浓度一效应关系的基础.
并推测两种浮萍对重金属的这种反应对植物适应环境变化有一定价值，也可以作为重
金属植物毒性评价的生物标记。重金属锅过量会对植物的生存形成多种胁迫效应，如
生长率的下降，光合作用效率降低，呼吸强度增加等，但它们能够促使稀脉萍和紫萍
群体发生解体的现象尚未见文献报道。
2.以铜、锡的胁迫效应为例，进一步研究了重金属产生的这种效应的机制，并测定了锅
胁迫下紫萍的一些生理生化指标的变化，较全面地了解了锅对紫萍的毒性效应。气相
色谱分析表明，在胁迫培养过程中，乙烯的产生量有很大增加，远远超过正常生长时
的产生量.说明在它们的胁迫下，有胁迫乙烯产生.根据已有的对其它植物进行研究
II
的报道，锅、铜胁迫能够刺激这些植物产生胁迫乙烯，并结合乙烯在植物体内具有的
能够加快衰老、促进脱叶的生理学作用，推侧?
Ecotoxicology is a newsubjecl just founded in about 1980 and is developing rapidlyOver the past two decades, ecotoxicology has experienced a rapid progress,thus it has become to be very important in protecting ecological safety from toxic contaminants. In our country, however, ecotoxicology is in its babyhood, to which we do not know very clear in many aspects. Enhancing the research in this field not only can faditate the advancement of this new subject in our country, but also greatly hasten the conversion of the economic pattern of our country into sustainable development.
Based on ecology, ecotoxicology is to study the hazardous influences of contaminants, especially, toxic chemicals on the structure and functions of ecosystem, the response of ecosystems to them, and the pathways of the toxicants moving, transforming and decomposing in the surrounding. Also, acotoxicology provides the technological base and tboretic steering for the ecological remediation of contaminated environment and ecological risk assessment. Of the toxic chemicals, heavy metals are given special attention by ecotocicological researchers because they are important inorganic poIlutantsEcotoxicology brought a profound revolution in toxicology, and led a change of the direction of the advance ofome environmentaltechniques.
Heavy metals have lots of effects on the plants, not only on the individuals, and on the populations and the comnunities in higher levels, but also in physiological and biochemical aspects and in genetics and evolution, e.g. in physiology, depressing the activity of some enzymes, initiating a few free radicals, reducing the growth of plants, degrading chlorophyll, inhibiting the elongation of the root and so on. But, the plant also develops plenty of mechanisms to resist the toxicities of heavy metals, such as stimulating stress proteins and stress ethylene production, activating antioxidant system to eliminate the fe radicals, synthesizing metallothioneins-like and phytochelatins, excluding more metals and accumulating and compartmentalizing in some organs. To understand the toxicity and mechanisms underlying of the heavy metals can increase our knowledge about thai ecological effects, and facilitate us to know their hazardous ecological effects more clear, which would urge us to use them
more availably and rationally so that their negative effects could be alleviated as possible.
Various chemicals, especially toxc chemicals, such as heavy metals or heavy metals-contained, insecticide, herbicide and so on, have great toxic effects orthe structure and functions ofinosphere. To assess their toxicity, including phytotoxicity, is the basis of scientific management fa them, and is profound to protect environmental quality and ecological safety.
Developed countries not only have weHestablished standard method to process the toxicity evaluation of toxicants, but also many related laws and rules for it. Our country drps greatly behind them in this aspect. However, the environmental status of our country has been so severe that we have to enhance the research in this field. It should be a short way to study the feasibility of using indigene organisms in assessing the taicity of pollutants based on using the established standard method. Management for reference, legislation and executing the law of toxic chemicals could be on the scientific basis when standard toxicity assessment method of our country is established. By far, the most commonly used vascular plants in toxicity tests are duckweeds, members of Lemanaceae.
This thesis conducted a completely ecotoxicological studyon the effects of heavy metals on two duckweed species, most common in our country,recollected from natural populations instead of from special strains selected in laboratory with axentic conditionsThe objects are to provide an available autochthonous model testing material for phytotoxicity assessment ofiiic chemicals in our country, an excellent plant material for phytoremediation of cadmimpolluted water. In
引文
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