苔藓植物对重金属污染的响应机理和生物指示的研究
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摘要
随着人类活动对环境干扰程度的日益加剧，利用生物对环境污染进行指示和检测成为当前对环境生物学研究的热点之一。苔藓植物作为一类良好的指示植物，被世界各国广泛应用为环境污染和变化的指示生物。但有关苔藓植物对环境污染的响应及机理研究仅有少数研究和报道。本论文在成功利用组培技术大量扩繁小立碗藓（Physcomitrella patens）、细叶小羽藓（Haplocladium microphyllum）和葡枝青藓（Brachythecium procumbens）配子体的基础上，采用先进的同步辐射X荧光分析技术（SRXRF）和生理生化测定分析，开展了组培条件下试管内苔藓植物的重金属胁迫实验，深入研究了重金属胁迫下3种藓类植物的重金属及营养元素的积累规律及其生理生化响应，探讨藓类植物对环境污染的响应机理及其生物指示作用。主要的研究结果如下：
（1）在大量的比较实验基础上，系统完成了细叶小羽藓和葡枝青藓的组培研究工作并建立了两种藓类配子体植株高效的人工快繁系统。研究发现，对于两种藓类植物配子体在试管内快繁而言，基本培养基的组成、培养基中外源蔗糖浓度、激素条件以及培养瓶内外状况都会对试管内生长的配子体产生重要影响。本研究最终确定的两种藓类配子体试管内快繁的最佳条件是：改良的Knop’s基本培养基添加10g/L的蔗糖；pH值调整在5.8至6.0；培养瓶内空气相对湿度保持在75%至85%之间；培养瓶要具有适度的透气性；培养温度为20±2℃；光照时间为14hr；光照强度为1500lux至2000lux。
上述相关研究成果获得了2项国家发明专利的授权，同时获得了重金属胁迫实验所需的两种藓类配子体材料。
（2）通过组培条件下的重金属胁迫实验，运用SRXRF技术，测定分析了不同浓度Pb~(2+)，Cu~(2+)，Cr~(6+)，Fe~(2+)等重金属离子单一或复合胁迫下，3种藓类的重金属元素的积累规律。单一重金属污染时，随着重金属离子污染浓度的增高，配子体中相应重金属元素的累积量呈现显著增高趋势。藓类植物种类不同，其配子体对重金属的累积能力和吸收敏感性也有差异。供试三种藓类比较，匐枝青藓植株体对Pb元素累积能力和吸收敏感性都非常强，其植株体中的Pb元素含量适合于作为评估环境Pb污染程度的量化指标。同样的比较分析表明，细叶小羽藓对Cr元素和Cu元素的累积能力和吸收敏感性都很强，其配子体内Cr元素和Cu元素的含量适合于作为这指示环境中两种重金属污染的量化指标。重金属复合胁迫对藓类配子体中重金属的累积能力有显著影响。以Fe~(2+)20mg/L与Pb~(2+)50mg/L、Cr~(6+)50mg/L、Cu~(2+)50mg/L形成的三种不同重金属复合污染处理时发现，较低浓度的Fe~(2+)对三种藓类吸收Pb~(2+)、Cr~(6+)、Cu~(2+)均具有强烈的拮抗作用，在很大程度上缓解了此三类重金属对藓类的毒害作用。采用u-SRXRF微探针扫描分析技术研究了配子体茎叶中重金属元素微区分布，在组织和细胞水平上揭示了藓类植物茎叶吸收累积重金属的机制。同时，进一步证实，供试三种藓类均具有强烈的重金属累积能力和敏感的生物响应特性，可作为监测环境重金属污染的生物指示植物。
（3）深入分析了重金属胁迫对藓类植物营养元素吸收的影响。结果表明，一定浓度的重金属离子对配子体生长所需的多种营养元素的吸收产生拮抗作用并抑制配子体对营养元素的吸收利用。在重金属单一污染条件下，随着污染浓度的增高，供试三种藓类植物配子体中营养元素的累积量呈现出两种变化趋势。第一种是营养元素累积量先升高后降低，即低污染浓度的重金属促进了藓类配子体中营养元素的累积，而过高污染浓度则会降低元素的累积量。第二种是营养元素累积量随污染浓度增高呈直接下降趋势。重金属种类不同，藓种不同，配子体中各种营养元素吸收和累积的受抑制强度不同。供试的Pb~(2+)，Cu~(2+)，Cr~(6+)，Fe~(2+)四种重金属离子污染下，细叶小羽藓配子体中P、K、Ca、S、Mn、Zn六种营养元素的最大降幅均低于小立碗藓，表明细叶小羽藓对上述4种重金属污染的耐受性都很强，最适合于作为环境重金属污染长期监测的指示植物。Pb~(2+)污染下，在最低污染浓度和最短污染时间的处理组，匐枝青藓配子体中P、K、Ca、S、Mn、Zn六种营养元素的含量就表现出显著的变化。说明匐枝青藓植株对铅污染产生的营养元素累积响应非常快速，适合作为铅污染的指示植物。在Pb~(2+)污染浓度100mg/L至Pb~(2+)污染浓度600mg/L之间，匐枝青藓配子体中P、K、Ca、S四种元素含量与Pb~(2+)污染浓度之间均呈现显著线性回归关系。说明在供试Pb~(2+)污染浓度范围内，Pb~(2+)污染浓度与匐枝青藓配子体中四种营养元素含量之间存在剂量-效应关系，建议利用匐枝青藓中的这四种营养元素含量作为监测环境重金属污染的辅助生理指标。不同种类重金属复合胁迫对于配子体不同种类营养元素吸收和累积表现出三种不同的胁迫效应，一种是缓解累积抑制效应，一种是增强累积抑制效应，一种是显著促进累积效应。
（4）采用生理生化分析方法，测定分析了不同重金属胁迫下3种藓类植物的生理响应和变化。结果表明，三种供试藓类总叶绿素含量对重金属污染的生物响应均非常快速，说明三种藓类植物对重金属污染反映敏感，适合于作为指示重金属污染的指示生物。Pb~(2+)污染导致三种藓类叶绿素b的降低幅度最大，说明叶绿素b含量的降低是Pb~(2+)污染导致植物光合作用受抑制的重要原因之一。不同种类单一重金属污染胁迫下，三种藓类植物POD酶活性响应表现出两种变化规律，一种是随着重金属污染浓度的增高，POD酶活性逐渐增强；一种是重金属低污染浓度范围内，POD酶活性逐渐增强，当污染浓度较高时，POD酶活性显著较低。供试三种藓类的POD酶活性与重金属污染浓度之间以及MDA含量与重金属污染浓度均存在显著的剂量-效应关系，可以拟合形成曲线回归关系显著的函数方程，因此建议将苔藓的POD酶活性和MDA含量作为监测环境重金属污染的辅助生理指标进行推广应用。
综上所述，本研究在分析3种藓类植物对重金属污染响应的基础上，进一步证明苔藓植物是作为环境重金属污染的敏感的良好的指示植物，为进一步利用苔藓植物监测和指示环境污染和变化提供了科学依据。
With the interference of human activities on the environment growing badly,biomonitoring of the polluted environments became one of hot points for study ofenvironmental biology. The bryophytes as good biological indicators to investigatepollution and changes of the environments have been widely used in differentcountries in the world. However, only few researches and reports dealing with theresponse and mechanism of the bryophytes to environmental pollution were carriedout. In order to better understanding the response and mechanism of bryophytes toenvironment pollution, the experiments of stress of different heavy metals on the threemosses, Haplocladium microphylum, Brachthecium procumbens and Physcomitriumpatens, which were successfully tissue cultivated and regenerated from native plants,were carried out on vitro condition at present study. Using advanced technique ofsynchrotron radiation x-fluorescence (SRXRF) and methods of physiological andbiochemical analyses, the accumulation of heavy metal and nutrient elements as wellas physiological and biochemical responses in the plants of three mosses under heavymetal stress were analyzed and studied. The main results are as follows:
(1) Based on numerous comparative experiments, the tissue cultivation andregeneration of the mosses Haplocladium microphyllum and Brachytheciumprocumbens were successfully completed and two kinds of systems of efficientartificial propagation of the gametophytes for the two mosses were established. Theresults showed the best conditions for cultivation of these two mosses are as follows:modified Knop's basic medium add10g/L sucrose; pH value adjusted at5.8to6.0;culture bottle relative air humidity kept at75%to85the%; culture bottles to have amoderate permeability; culture temperature was20±2℃; illumination time was14hr; the illumination the strength for1500lux to2000lux. As the results, twoinvention patents were applied and obtained, and plants material for the experiments of heavy metal stress was gathered.
(2) In the stress experiments under vitro condition, the different concentrationsof heavy metals ions including Pb, Cu, Cr, Fe in the plants of three mosses under thestress of single or combined condition of different heavy metals were analyzed bySRSRF. Under the stress of single heavy metal element, with increase of the pollutionconcentration, the cumulative amount of the same heavy metal ions had higher trendof significantly increasing in the plants. But, accumulation capacity and absorptionsensitivity of the heavy metals were different among three mosses. In comparison ofthe tested three mosses, Brachythecium procumbens had the highest content of Pb of1699.05147.14μg/g and152,9813.22μg/g in the plants under the highest Pb~(2+)400mg/L pollution and lowest Pb~(2+)50mg/L pollution, respectively. It indicated thatcumulative capacity and absorption sensitivity of Pb for Brachythecium procumbensare very strong and quantitative indicator of the Pb content in the plant is suitable forthe assessment of environmental Pb pollution levels. Same analysis showed thatHaplocladium microphyllum also had strong cumulative capacity and absorptionsensitivity of the Cr and Cu, so the content of the Cr and Cu elements in the plants issuitable indicators for Cr and Cu pollution. The combined stress of different elementssignificantly impact of the accumulation of heavy metals in the moss plants. Underthe three combined treatments of Fe~(2+)20mg/L with Pb~(2+)50mg/L, Cr~(6+)50mg/L,Cu~(2+)50mg/L, while Fe cumulative amount significantly increased, cumulativeamounts of Pb, Cr, Cu in the plants are significantly lower than those under the stressof single treatment with same element. This showed that the lower tconcentration ofFe~(2+)had a strong sense of antagonism on Pb~(2+), Cr~(6+), Cu~(2+)and can largely alleviatepoisoning effect of three heavy metals on the mosses. In addition, the scanningtechnology of U-SRXRF micro-probe analysis was used to show micro-distribution ofheavy metals on the stem and leaves of the mosses. The study reveals absorptionmechanism of accumulation of heavy metals in the tissue and cell levels in the mosses.At the same time, it is further confirmed that the tested three mosses have a strongaccumulation of heavy metals and sensitive biological response and can be used asindicator plants for biological monitoring of environmental heavy metal pollution.
(3) The absorption capacity and changes of different nutrient elements including,P、K、Ca、S、Mn、Zn under different heavy metal stress were also analyzed. The resultsshowed that a certain concentration of heavy metal ions could have antagonistic effect on the absorption of the nutrient elements and inhibit absorption and utilization ofnutrient elements in the plants. With the increasing concentration of single heavymetal pollution, nutritive elements cumulative amount of the tested three mossesshowed two trends. The first one is that lower concentration of heavy metal promotedthe absorption of nutritional elements while excessive pollution concentrationsreduced the cumulative amount of the element. The second is the amount of nutrientaccumulation was directly downward with the increased pollution concentrations. Theabsorption and accumulation of nutrients were effected by different elements as wellas different mosses. Under stress treatment of Pb~(2+), Cu~(2+), Cr~(2+), Fe~(2+)the lowestdecrease of P, K, Ca, S, Mn, and Zn nutrient elements in Haplocladium microphyllumwere high than in Physcomitrella patens. It implied that Haplocladium microphyllumhas strong tolerance to the four heavy metals pollution and is the most suitable forlong-term monitoring of the environmental heavy metal pollution. The results that thesignificant changes of P, K, Ca, S, Mn, and Zn nutrient elements in Brachytheciumprocumbens under the lowest concentration and shortest time treatment of Pb~(2+)alsoindicated that Brachythecium procumbens responses to Pb pollution rapidly and canbe used as indicative plant to Pb pollution. Linear regression analysis of the resultsshowed that, between Pb~(2+)pollution concentration of100mg/L to Pb~(2+)pollutionconcentration of600mg/L, the content of P, K, Ca, S in the plants of Brachytheciumprocumbens and Pb~(2+)pollution concentration presented a significant line regression. Itis recommended that the use of nutritional element content in the plants ofBrachythecium procumbens as additional indicators for monitoring of environmentalPb pollution.The impact of different kinds of combined heavy metals stress ondifferent mosses on nutrient absorption and accumulation capacity is obviouslydifferent. The impact could be divided into three types: a remission cumulativeinhibitory effect; enhanced cumulative inhibitory effect, and a significant promotionof cumulative effect.
(4) The physiological response and changes of three species of mosses includingphotosynthetic pigment synthesis, peroxidase (POD) activity changes,malondialdehyde (MDA) content under heavy metal stress were measured andcompared. The fact that the biological responses of total chlorophyll content to heavymetal contamination in the mosses were very rapid indicated that the mosses aresensitive and suitable for monitoring heavy metal pollution. Pb~(2+)pollution cause thethree mosses chlorophyll b decrease. The largest decline of chlorophyll b of Physcomitrella patens, Haplocladium microphyllum and Brachythecium procumbenswere58.55%,48.33%and46.17%, respectively. It implied that one of importantreasons of inhibiting plant photosynthesis is lower content of chlorophyll b caused byPb~(2+)pollution. Under the different types of single heavy metal pollution stress, PODactivity response of three mosses showed two kinds of variation, a POD enzymeactivity gradually increased with increasing concentration of heavy metal pollution;and POD enzyme activity gradually enhanced in a concentration range of heavymetals polluting and then significantly lower under higher pollution concentrations.Significant dose-response relationship between POD activity and MDA content andconcentration of heavy metal pollution in the test three mosses fitted to the formationof significant functional equation of the curve regression relationship. Therefore, it isrecommended that the moss POD enzyme activity and MDA content can be used asauxiliary physiological indicators for monitoring environmental heavy metalpollution.
In summary, the present study on the response analysis of three mosses to heavymetal pollution provided further evidence that bryophytes are sensitive and goodindicator plants to environmental pollution of heavy metals. The research resultsprovided scientific basis for further study and application of monitoringenvironmental pollution and changes by bryophytes.
（1）在大量的比较实验基础上，系统完成了细叶小羽藓和葡枝青藓的组培研究工作并建立了两种藓类配子体植株高效的人工快繁系统。研究发现，对于两种藓类植物配子体在试管内快繁而言，基本培养基的组成、培养基中外源蔗糖浓度、激素条件以及培养瓶内外状况都会对试管内生长的配子体产生重要影响。本研究最终确定的两种藓类配子体试管内快繁的最佳条件是：改良的Knop’s基本培养基添加10g/L的蔗糖；pH值调整在5.8至6.0；培养瓶内空气相对湿度保持在75%至85%之间；培养瓶要具有适度的透气性；培养温度为20±2℃；光照时间为14hr；光照强度为1500lux至2000lux。
上述相关研究成果获得了2项国家发明专利的授权，同时获得了重金属胁迫实验所需的两种藓类配子体材料。
（2）通过组培条件下的重金属胁迫实验，运用SRXRF技术，测定分析了不同浓度Pb~(2+)，Cu~(2+)，Cr~(6+)，Fe~(2+)等重金属离子单一或复合胁迫下，3种藓类的重金属元素的积累规律。单一重金属污染时，随着重金属离子污染浓度的增高，配子体中相应重金属元素的累积量呈现显著增高趋势。藓类植物种类不同，其配子体对重金属的累积能力和吸收敏感性也有差异。供试三种藓类比较，匐枝青藓植株体对Pb元素累积能力和吸收敏感性都非常强，其植株体中的Pb元素含量适合于作为评估环境Pb污染程度的量化指标。同样的比较分析表明，细叶小羽藓对Cr元素和Cu元素的累积能力和吸收敏感性都很强，其配子体内Cr元素和Cu元素的含量适合于作为这指示环境中两种重金属污染的量化指标。重金属复合胁迫对藓类配子体中重金属的累积能力有显著影响。以Fe~(2+)20mg/L与Pb~(2+)50mg/L、Cr~(6+)50mg/L、Cu~(2+)50mg/L形成的三种不同重金属复合污染处理时发现，较低浓度的Fe~(2+)对三种藓类吸收Pb~(2+)、Cr~(6+)、Cu~(2+)均具有强烈的拮抗作用，在很大程度上缓解了此三类重金属对藓类的毒害作用。采用u-SRXRF微探针扫描分析技术研究了配子体茎叶中重金属元素微区分布，在组织和细胞水平上揭示了藓类植物茎叶吸收累积重金属的机制。同时，进一步证实，供试三种藓类均具有强烈的重金属累积能力和敏感的生物响应特性，可作为监测环境重金属污染的生物指示植物。
（3）深入分析了重金属胁迫对藓类植物营养元素吸收的影响。结果表明，一定浓度的重金属离子对配子体生长所需的多种营养元素的吸收产生拮抗作用并抑制配子体对营养元素的吸收利用。在重金属单一污染条件下，随着污染浓度的增高，供试三种藓类植物配子体中营养元素的累积量呈现出两种变化趋势。第一种是营养元素累积量先升高后降低，即低污染浓度的重金属促进了藓类配子体中营养元素的累积，而过高污染浓度则会降低元素的累积量。第二种是营养元素累积量随污染浓度增高呈直接下降趋势。重金属种类不同，藓种不同，配子体中各种营养元素吸收和累积的受抑制强度不同。供试的Pb~(2+)，Cu~(2+)，Cr~(6+)，Fe~(2+)四种重金属离子污染下，细叶小羽藓配子体中P、K、Ca、S、Mn、Zn六种营养元素的最大降幅均低于小立碗藓，表明细叶小羽藓对上述4种重金属污染的耐受性都很强，最适合于作为环境重金属污染长期监测的指示植物。Pb~(2+)污染下，在最低污染浓度和最短污染时间的处理组，匐枝青藓配子体中P、K、Ca、S、Mn、Zn六种营养元素的含量就表现出显著的变化。说明匐枝青藓植株对铅污染产生的营养元素累积响应非常快速，适合作为铅污染的指示植物。在Pb~(2+)污染浓度100mg/L至Pb~(2+)污染浓度600mg/L之间，匐枝青藓配子体中P、K、Ca、S四种元素含量与Pb~(2+)污染浓度之间均呈现显著线性回归关系。说明在供试Pb~(2+)污染浓度范围内，Pb~(2+)污染浓度与匐枝青藓配子体中四种营养元素含量之间存在剂量-效应关系，建议利用匐枝青藓中的这四种营养元素含量作为监测环境重金属污染的辅助生理指标。不同种类重金属复合胁迫对于配子体不同种类营养元素吸收和累积表现出三种不同的胁迫效应，一种是缓解累积抑制效应，一种是增强累积抑制效应，一种是显著促进累积效应。
（4）采用生理生化分析方法，测定分析了不同重金属胁迫下3种藓类植物的生理响应和变化。结果表明，三种供试藓类总叶绿素含量对重金属污染的生物响应均非常快速，说明三种藓类植物对重金属污染反映敏感，适合于作为指示重金属污染的指示生物。Pb~(2+)污染导致三种藓类叶绿素b的降低幅度最大，说明叶绿素b含量的降低是Pb~(2+)污染导致植物光合作用受抑制的重要原因之一。不同种类单一重金属污染胁迫下，三种藓类植物POD酶活性响应表现出两种变化规律，一种是随着重金属污染浓度的增高，POD酶活性逐渐增强；一种是重金属低污染浓度范围内，POD酶活性逐渐增强，当污染浓度较高时，POD酶活性显著较低。供试三种藓类的POD酶活性与重金属污染浓度之间以及MDA含量与重金属污染浓度均存在显著的剂量-效应关系，可以拟合形成曲线回归关系显著的函数方程，因此建议将苔藓的POD酶活性和MDA含量作为监测环境重金属污染的辅助生理指标进行推广应用。
综上所述，本研究在分析3种藓类植物对重金属污染响应的基础上，进一步证明苔藓植物是作为环境重金属污染的敏感的良好的指示植物，为进一步利用苔藓植物监测和指示环境污染和变化提供了科学依据。
With the interference of human activities on the environment growing badly,biomonitoring of the polluted environments became one of hot points for study ofenvironmental biology. The bryophytes as good biological indicators to investigatepollution and changes of the environments have been widely used in differentcountries in the world. However, only few researches and reports dealing with theresponse and mechanism of the bryophytes to environmental pollution were carriedout. In order to better understanding the response and mechanism of bryophytes toenvironment pollution, the experiments of stress of different heavy metals on the threemosses, Haplocladium microphylum, Brachthecium procumbens and Physcomitriumpatens, which were successfully tissue cultivated and regenerated from native plants,were carried out on vitro condition at present study. Using advanced technique ofsynchrotron radiation x-fluorescence (SRXRF) and methods of physiological andbiochemical analyses, the accumulation of heavy metal and nutrient elements as wellas physiological and biochemical responses in the plants of three mosses under heavymetal stress were analyzed and studied. The main results are as follows:
(1) Based on numerous comparative experiments, the tissue cultivation andregeneration of the mosses Haplocladium microphyllum and Brachytheciumprocumbens were successfully completed and two kinds of systems of efficientartificial propagation of the gametophytes for the two mosses were established. Theresults showed the best conditions for cultivation of these two mosses are as follows:modified Knop's basic medium add10g/L sucrose; pH value adjusted at5.8to6.0;culture bottle relative air humidity kept at75%to85the%; culture bottles to have amoderate permeability; culture temperature was20±2℃; illumination time was14hr; the illumination the strength for1500lux to2000lux. As the results, twoinvention patents were applied and obtained, and plants material for the experiments of heavy metal stress was gathered.
(2) In the stress experiments under vitro condition, the different concentrationsof heavy metals ions including Pb, Cu, Cr, Fe in the plants of three mosses under thestress of single or combined condition of different heavy metals were analyzed bySRSRF. Under the stress of single heavy metal element, with increase of the pollutionconcentration, the cumulative amount of the same heavy metal ions had higher trendof significantly increasing in the plants. But, accumulation capacity and absorptionsensitivity of the heavy metals were different among three mosses. In comparison ofthe tested three mosses, Brachythecium procumbens had the highest content of Pb of1699.05147.14μg/g and152,9813.22μg/g in the plants under the highest Pb~(2+)400mg/L pollution and lowest Pb~(2+)50mg/L pollution, respectively. It indicated thatcumulative capacity and absorption sensitivity of Pb for Brachythecium procumbensare very strong and quantitative indicator of the Pb content in the plant is suitable forthe assessment of environmental Pb pollution levels. Same analysis showed thatHaplocladium microphyllum also had strong cumulative capacity and absorptionsensitivity of the Cr and Cu, so the content of the Cr and Cu elements in the plants issuitable indicators for Cr and Cu pollution. The combined stress of different elementssignificantly impact of the accumulation of heavy metals in the moss plants. Underthe three combined treatments of Fe~(2+)20mg/L with Pb~(2+)50mg/L, Cr~(6+)50mg/L,Cu~(2+)50mg/L, while Fe cumulative amount significantly increased, cumulativeamounts of Pb, Cr, Cu in the plants are significantly lower than those under the stressof single treatment with same element. This showed that the lower tconcentration ofFe~(2+)had a strong sense of antagonism on Pb~(2+), Cr~(6+), Cu~(2+)and can largely alleviatepoisoning effect of three heavy metals on the mosses. In addition, the scanningtechnology of U-SRXRF micro-probe analysis was used to show micro-distribution ofheavy metals on the stem and leaves of the mosses. The study reveals absorptionmechanism of accumulation of heavy metals in the tissue and cell levels in the mosses.At the same time, it is further confirmed that the tested three mosses have a strongaccumulation of heavy metals and sensitive biological response and can be used asindicator plants for biological monitoring of environmental heavy metal pollution.
(3) The absorption capacity and changes of different nutrient elements including,P、K、Ca、S、Mn、Zn under different heavy metal stress were also analyzed. The resultsshowed that a certain concentration of heavy metal ions could have antagonistic effect on the absorption of the nutrient elements and inhibit absorption and utilization ofnutrient elements in the plants. With the increasing concentration of single heavymetal pollution, nutritive elements cumulative amount of the tested three mossesshowed two trends. The first one is that lower concentration of heavy metal promotedthe absorption of nutritional elements while excessive pollution concentrationsreduced the cumulative amount of the element. The second is the amount of nutrientaccumulation was directly downward with the increased pollution concentrations. Theabsorption and accumulation of nutrients were effected by different elements as wellas different mosses. Under stress treatment of Pb~(2+), Cu~(2+), Cr~(2+), Fe~(2+)the lowestdecrease of P, K, Ca, S, Mn, and Zn nutrient elements in Haplocladium microphyllumwere high than in Physcomitrella patens. It implied that Haplocladium microphyllumhas strong tolerance to the four heavy metals pollution and is the most suitable forlong-term monitoring of the environmental heavy metal pollution. The results that thesignificant changes of P, K, Ca, S, Mn, and Zn nutrient elements in Brachytheciumprocumbens under the lowest concentration and shortest time treatment of Pb~(2+)alsoindicated that Brachythecium procumbens responses to Pb pollution rapidly and canbe used as indicative plant to Pb pollution. Linear regression analysis of the resultsshowed that, between Pb~(2+)pollution concentration of100mg/L to Pb~(2+)pollutionconcentration of600mg/L, the content of P, K, Ca, S in the plants of Brachytheciumprocumbens and Pb~(2+)pollution concentration presented a significant line regression. Itis recommended that the use of nutritional element content in the plants ofBrachythecium procumbens as additional indicators for monitoring of environmentalPb pollution.The impact of different kinds of combined heavy metals stress ondifferent mosses on nutrient absorption and accumulation capacity is obviouslydifferent. The impact could be divided into three types: a remission cumulativeinhibitory effect; enhanced cumulative inhibitory effect, and a significant promotionof cumulative effect.
(4) The physiological response and changes of three species of mosses includingphotosynthetic pigment synthesis, peroxidase (POD) activity changes,malondialdehyde (MDA) content under heavy metal stress were measured andcompared. The fact that the biological responses of total chlorophyll content to heavymetal contamination in the mosses were very rapid indicated that the mosses aresensitive and suitable for monitoring heavy metal pollution. Pb~(2+)pollution cause thethree mosses chlorophyll b decrease. The largest decline of chlorophyll b of Physcomitrella patens, Haplocladium microphyllum and Brachythecium procumbenswere58.55%,48.33%and46.17%, respectively. It implied that one of importantreasons of inhibiting plant photosynthesis is lower content of chlorophyll b caused byPb~(2+)pollution. Under the different types of single heavy metal pollution stress, PODactivity response of three mosses showed two kinds of variation, a POD enzymeactivity gradually increased with increasing concentration of heavy metal pollution;and POD enzyme activity gradually enhanced in a concentration range of heavymetals polluting and then significantly lower under higher pollution concentrations.Significant dose-response relationship between POD activity and MDA content andconcentration of heavy metal pollution in the test three mosses fitted to the formationof significant functional equation of the curve regression relationship. Therefore, it isrecommended that the moss POD enzyme activity and MDA content can be used asauxiliary physiological indicators for monitoring environmental heavy metalpollution.
In summary, the present study on the response analysis of three mosses to heavymetal pollution provided further evidence that bryophytes are sensitive and goodindicator plants to environmental pollution of heavy metals. The research resultsprovided scientific basis for further study and application of monitoringenvironmental pollution and changes by bryophytes.
引文
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