不同条件下浮萍磷吸收效率及其作用机理
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摘要
随着经济社会发展,水体富营养化问题日趋严重,其中以磷的贡献尤为突出。如何经济、有效地去除磷污染是解决水体富营养化问题的关键。近年来,利用特定水生植物进行水体生态修复的研究引起了广泛关注。研究表明,浮萍除磷具有效率高、成本低的优点。目前,关于浮萍净化污水的研究主要集中于浮萍的生长繁殖能力,以及对氮的吸收机理研究。但涉及影响浮萍磷吸收的因素及其机理的研究仍较为有限。本研究围绕不同因子对浮萍磷吸收效率的影响,通过对浮萍磷吸收效率、磷吸收动力学特征、与磷代谢相关的酶活性以及细胞微结构变化的研究,详细阐述了品种差异、光照强度、温度、pH、水体磷浓度以及不同类型富营养化水体环境对浮萍磷吸收效率的影响,明确了不同条件下浮萍的磷吸收效率,阐明了其作用机理,为提高浮萍的磷净化效率提供了理论依据。通过研究得出以下主要结果：
(1)稀脉浮萍的相对生长率RGR高于少根紫萍和紫萍,稀脉浮萍叶状体增殖速度最快。在实验室模拟培养条件下,稀脉浮萍叶状体的倍增时间最短,为3.23,少根紫萍和紫萍叶状体倍增时间分别为4.15天和4.81天。单位鲜重稀脉浮萍吸收的磷量显著低于少根紫萍和紫萍,其高磷累积吸收量建立在生物量大量扩增基础上。稀脉浮萍对H2PO4-的的亲和力常数Km和最大吸收速率Vmax均最小,紫萍的Km和Vmax最大。稀脉浮萍对H2PO4-的亲和力更强,在较低的离子浓度下就可以达到Vmax；而当环境中磷浓度较高时,紫萍对磷的净化速率高于少根紫萍和稀脉浮萍。实验室模拟培养期间,紫萍体内的ACPase和H+-ATPase活力水平始终最高,稀脉浮萍次之,少根紫萍最低。从细胞学形态结构上分析,紫萍和少根紫萍叶绿体基粒内囊体片层结构排列紧密,叶绿体周围环绕多个线粒体,而稀脉浮萍叶绿体中存在较大的淀粉粒构和空泡结构,片层结构松散。紫萍和少根紫萍的细胞超微结构更有利于植物光能利用效率和同化物运输能力的提高。因此,紫萍对磷的吸收能力更强,更适于对高磷浓度污水进行修复。
(2)紫萍RGR随光照强度增加而逐渐上升,在60001x高光强下,紫萍叶状体仍维持较高扩增速度。紫萍累积吸收磷量随着光照强度提高而升高,但单位鲜重浮萍吸收的磷量在60001x高光强条件下最低,30001x光强时的磷吸收能力略高于15001x,但两者之间无显著差异。随着光照强度增加,紫萍对H2P04-的最大吸收速率Vmax增加；不同光强条件下紫萍对H2P04-的亲和力常数Km略有差异,3000lx和60001x光强下略高于15001x光强。随着光照强度的增加,紫萍体内ACPase活力增加。实验室模拟培养期间,紫萍H+-ATP酶活力呈先缓慢下降随后逐渐上升的现象。培养后期低光强下,紫萍的H+-ATP酶活力超过高光强处理。强光下,紫萍叶肉细胞内的叶绿体数明显增加；弱光时虽然叶绿体数较少,但其基粒片层结构变厚,增加了光合作用反应面积。
(3)25℃时紫萍RGR最高,35℃略低,10℃最低。低温对紫萍生长的胁迫作用甚于高温。紫萍的累积吸收磷量随着温度的升高而增加,但单位鲜重浮萍吸收的磷量在10℃时最高,35℃次之,25℃最低。25℃和35℃条件下,紫萍的高磷累积吸收量通过生物量的大量扩增实现。随着温度增加,紫萍对H2P04-的最大吸收速率Vmax增加：不同温度之间亲和力大小依次为：25℃>10℃>35℃。35℃条件下紫萍对H2PO4-的低亲和力可能是高温胁迫作用所致。随着温度增加,紫萍体内ACPase活力增加。但长时间的低温胁迫也会使紫萍体内ACP酶活力显著提高。不同温度条件下,紫萍体内H+-ATPase活力的大小依次为：25℃>10℃。从细胞超微结构来看,与25℃适温条件相比,10℃低温和35℃高温下,紫萍细胞均存在一定程度的逆境胁迫作用。表现为低温时部分叶绿体皱缩成带状,叶肉细胞之间出现较大空隙；高温时叶绿体外膜溶解,基质外渗,脂质小球数量增多。
(4)不同pH条件下,紫萍RGR大小依次为pH6>pH5>pH7>pH8>pH9。紫萍磷累积吸收量在pH6条件下最高,当pH>6时,随着pH升高,磷累积吸收量降低。pH对单位鲜重紫萍吸收的磷量无显著影响。不同pH条件下,紫萍对H2P04-的最大吸收速率Vmax大小依次为：pH6>pH7>pH8>pH5>pH9,亲和力常数Km大小依次为：pH5>pH9>pH8>pH7>pH6。不同pH条件培养较长时间后,紫萍H+-ATPase和ACPase活力在pH6、pH7和pH8条件下高于pH5和pH9。紫萍在pH5条件下叶肉细胞中的叶绿体数量明显减少,类囊体片层结构松散。在pH6和pH7条件下,紫萍叶绿体数量丰富,叶肉细胞中溶质和细胞器较多,类囊体片层排列整齐紧密,光合效率高。pH8和pH9条件下,紫萍植株受到高OH-伤害,尤其是在pH9时,叶肉细胞中的叶绿体数量急剧减少,叶绿体和类囊体膜崩解,形成大量脂质小球。
(5)紫萍在水体总磷浓度0.1mg/L～15mg/L范围内,能保持较高的RGR,累积吸收磷量和单位鲜重紫萍吸收磷量均随着磷浓度升高而增加,当水体中总磷浓度达到45mg/L以上时,紫萍生长受限,RGR较低,磷累积吸收量和单位鲜重紫萍吸收的磷量均显著下降。当水体中总磷浓度在0.3mg/L-3mg/L范围时,紫萍对水体中磷的去除效率最高(约70%)。当水体中总磷浓度低于0.3mg/L时,紫萍体内ACPase活力升高,但长时间的低磷条件使紫萍生长受抑,从而使低磷和高磷条件之间ACPase活力的差异减小。与高磷条件相比,低磷条件下紫萍H+-ATPase活力较低。当水体中磷浓度较低时,紫萍叶绿体中出现大量淀粉粒结构,光合产物运输受阻；提高水体环境中磷浓度,叶绿体中淀粉粒结构开始消失；当进一步提高磷浓度使水体中总磷浓度达到45mg/L时,紫萍细胞出现质壁分离现象,叶绿体结构受到破坏,表现为外膜溶解,类囊体片层排列紊乱,对营养物的同化效率降低。
(6)不同类型富营养化水体中,三种浮萍的磷吸收效率存在差异。稀脉浮萍在氮磷浓度较低的景观水体中,磷累积吸收量最高。由于磷累积吸收量与生物量几乎按比例同步增加,以鲜重为基础算得的单位鲜重浮萍吸收的磷量并没有增加。紫萍在氮磷浓度较高的农业废水和生活污水中,磷累积吸收量和单位鲜重浮萍吸收的磷量均高于少根紫萍和稀脉浮萍,表现出对高磷浓度水体的磷吸收优势。稀脉浮萍叶绿素含量在景观水体中最高,在生活污水中最低。少根紫萍和紫萍叶绿素含量始终随着水体中氮磷浓度上升而升高。三种浮萍蛋白质含量均随着水体中氮磷浓度的升高而升高。不同类型富营养化水体中,稀脉浮萍体内蛋白质含量始终高于少根紫萍和紫萍。紫萍体内ACPase和H+-ATPase活力随着水体中磷浓度的增加而升高,在磷浓度较高的农业废水和生活污水中紫萍的ACPase和H+-ATPase活力显著高于稀脉浮萍和少根紫萍。在高磷浓度水体中,紫萍对污水中磷的去除效果、生长速率以及对磷的吸收积累能力均高于低磷环境,适于修复高磷污染的生活污水和农业废水；稀脉浮萍在低磷浓度水体中的生长速率高、对磷的吸收能力强,适于低磷污染的景观水体修复。
(7)用Hill方程对三种品种浮萍吸收H2P04速率与溶液中磷浓度关系进行拟合,其拟合度都高于Michaelis-Menten方程。不同环境条件下(包括光照强度、温度、pH、水体中初始磷浓度水平),对比Hill方程和Michaelis-Menten方程拟合的紫萍吸收HPO42-的速率与溶液中总磷浓度的关系曲线发现,前者的拟合效果均不同程度上优于后者。说明在较宽的磷浓度范围内,浮萍对H2P04的吸收速率与溶液中磷浓度之间表现出“S形”吸收曲线关系。
With the fast development of economy and population, water body eutrophication has become one of the most important environmental problems. Phosphorus is known to play an important role in water eutrophication. In order to improve water quality it is crucial to remove phosphorus from eutrophic water effectively. In recent years, ecological remediation methods using aquatic plants have aroused widespread interest. One of the most efficient and economical ways is the use of duckweed for eutrophic water body treatment. However, studies to date mainly deal with duckweed growth capacity and mechanisms of nitrogen uptake, while knowledge concerning mechanisms of phosphorus uptake and controlling factors still remains limited. In this paper, phosphorus purification capacity of duckweed were studied by determining phosphorus uptake efficiency, uptake kinetics, characteristics of phosphorus metabolism enzymes in plants and cell ultrastructure changes under the influence of various factors(varieties, light intensity, temperature, pH, phosphorus concentration and water type). The absorption efficiency and mechanisms of phosphorus by duckweed under different conditions were determined, which provided an important theoretical reference for the development and utilization of duckweed in the phosphorus restoration of eutrophic water bodies. The main results were as follows:
(1) Lemna aequinoctialis had a higher relative growth rate than Spirodela oligorrhiza and Spirodela polyrrhiza, with a leaflets'doubling time of3.23days, while the doubling times of Spirodela oligorrhiza and Spirodela polyrrhiza were4.15and4.81days respectively. However, the phosphorus uptake capacity of Lemna aequinoctialis was siginificantly lower than that of Spirodela oligorrhiza and Spirodela polyrrhiza. The high phosphorus accumulation in Lemna aequinoctialis was a result of biomass increase. The Km and Vmax values for phosphorus uptake of Lemna aequinoctialis were lower than those of Spirodela oligorrhiza and Spirodela polyrrhiza, indicating a stronger affinity to phosphorus of Lemna aequinoctialis Spirodela polyrrhiza had the highest Km and Vmax values for phosphorus uptake, which might lead to a better performance of Spirodela polyrrhiza in purifying wastewater with a higher phosphorus concentration. Spirodela polyrrhiza had the highest ACPase and H+-ATPase activity, followed by Lemna aequinoctialis and Spirodela oligorrhiza. Morphologically, Spirodela oligorrhiza and Spirodela polyrrhiza had tightly arranged chloroplasts thylakoid membranes, and the chloroplasts were surrounded by many mitochondrias, while chloroplasts in Lemna aequinoctialis had relatively loose structure, with accumulated starch and vesicles. Cell ultrastructure of Spirodela oligorrhiza and Spirodela polyrrhiza was of great benefit to light use and transport of assimilation substance in plants. Therefore, Spirodela polyrrhiza had higher phosphorus absorption capability than Spirodela oligorrhiza and Lemna aequinoctialis, making it more suitable for purifying high-phosphorus polluted water.
(2) RGR value of Spirodela polyrrhiza increased with the increase of light intensity. Spirodela polyrrhiza fronds remained in a high expanding speed even in60001x high light intensity. Total absorbed phosphorus amount of Spirodela polyrrhiza increased with the increase of light intensity, while phosphorus uptake capacity was the lowest at60001x high light intensity. There was no significant difference between phosphorus uptake capacity with light intensity of30001x and15001x. With the increase of light intensity Vmax for phosphorus uptake of Spirodela polyrrhiza increased, while no change was observed for Km. Phosphorus affinity of Spirodela polyrrhiza with30001x and60001x light intensity were slightly higher than15001x. ACPase activity of Spirodela polyrrhiza also increased with the increase of light intensity. H+-ATPase activity firstly decreased, and then increased slowly during the cultivation period. At the later period of cultivation, H+-ATPase activity of Spirodela polyrrhiza with weak light intensity were higher than strong light intensity. When supplied with strong light intensity, chloroplasts in mesophyll cell of Spirodela polyrrhiza increased. Though the number of chloroplasts decreased with weak light intensity, grana lamella of chloroplasts thickened to increase its photosynthesis reaction area.
(3) The highest and lowest RGR values of Spirodela polyrrhiza were determined at25℃and10℃respectively. The growth of duckweed was negatively influenced more by low temperature than by high temperature. Total absorbed phosphorus in Spirodela polyrrhiza increased with temperature, while the highest phosphorus uptake capacity was detected at10℃, which indicated that biomass increase was the reason for high cumulation of absorbed phosphorus in25℃and35℃treatments. Vmax for phosphorus uptake of Spirodela polyrrhiza increased with the increase of temperature. Km value of phosphorus affinity changed at different temperatures:25℃>10℃>35℃. ACPase activity of Spirodela polyrrhiza increased with the increase of temperature, however, the stressing effect caused by a long period of time with low temperature may also lead to a higher phosphorus affinity. H+-ATPase activity of Spirodela polyrrhiza under different temperature was25℃>10℃>35℃. In terms of cell ultrastructure, Spirodela polyrrhiza was under stress to a certain degree in10℃and35℃treatments, as shown by shriveled. Chloroplasts and large cell cavity at10℃, and dissolved chloroplast envelope, exuded stroma and increased Plastoglobulus at35℃.
(4) RGR values of Spirodela polyrrhiza at different pH were pH6> pH5> pH7> pH8> pH9. The highest total cumulative absorbed phosphorus in Spirodela polyrrhiza was found at pH6, while phosphorus uptake capacity was not affected by pH. Vmax for phosphorus uptake of Spirodela polyrrhiza was pH6> pH7> pH8>pH5> pH9, and Km was pH5> pH9> pH8> pH7> pH6. H+-ATPase and ACPase activities of Spirodela polyrrhiza were higher at pH6, pH7and pH8than other treatments. The number of chloroplasts in mesophyll cell of Spirodela polyrrhiza decreased significantly at pH5, with loose strucutre of thylakoidas lamella. At pH6and pH7, Spirodela polyrrhiza had more chloroplasts, solvend and organelles, and tightly arranged thylakoids, which assured a high photosynthetic efficiency. Duckweed was damaged by high concentration of OH-under pH8and pH9treatments, especially at pH9. As a result, the number of chloroplasts decreased dramatically, while the membranes of chloroplasts and thylakoid were collapsed and many plastoglobulus ermerged.
(5) Spirodela polyrrhiza could maintain a high RGR in water with0.1mg/L to15mg/L of phosphorus. The growth of Spirodela polyrrhiza was restricted, and RGR value was low as phosphorus concentration in water was above45mg/L. The highest phosphorus removal efficiency of Spirodela polyrrhiza was reached approximately70%when phosphorus concentration in solution was ranged between0.3mg/L and3mg/L. ACPase activity of Spirodela polyrrhiza increased when phosphorus was lower than0.3mg/L. However, long time exposure to low phosphorus would limit the growth of Spirodela polyrrhiza, which thus reduced the difference of ACPase activities betweenlow and high phosphorus concentration treatments. H+-ATPase activity with high phosphorus treatment was higher than low phosphorus treatment. Under low phosphorus treatment, many starch grains appeared in chloroplasts and the transport of photosynthetic products was interrupted. As soon as phosphorus concentration of outer solution was increased, starch grains disappeared. When phosphorus concentration reached45mg/L, plasm olysis started and structure of chloroplasts was destroyed, indicated by disintegrated membranes, lamella of thylakoidas arranged in disorder, and decreased nutrients assimilatory efficiency.
(6) Difference existed in phosphorus uptake efficiency of three duckweek species in different types of eutrophic water. In landscape water with low nitrogen and phosphorus concentrations, total absorbed phosphorus in Lemna aequinoctialis was higher than in Spirodela oligorrhiza and Spirodela polyrrhiza. This is due to the increase of biomass. However, total absorbed phosphorus and phosphorus uptake capacity of Spirodela polyrrhiza was the highest in domestic sewage and agricultural waste water with higher nitrogen and phosphorus concentration. Spirodela polyrrhiza showed a high phosphorus uptake capacity in high phosphorus concentration environment. In landscape water, chlorophyll content of Lemna aequinoctialis was higher than the other two species, whereas in domestic sewage it was the lowest. With the increase of nitrogen and phosphorus concentration in water, chlorophyll content of Spirodela oligorrhiza and Spirodela polyrrhiza increasedand protein content also increasedin all three duckweed species. ACPase and H+-ATPase activity of Spirodela polyrrhiza, which increased with the increase of phosphorus concentration of outer water environment, was much higher than that of Lemna aequinoctialis and Spirodela oligorrhiza in domestic sewage and agricultural waste water treatments. In high phosphorus water environment, phosphorus removal efficiency, cumulative uptake capacity, and growth rate of Spirodela polyrrhiza were higher than low phosphorus environment, making it a better candidate for treating domestic sewage and agricultural waste water with a high phosphorus concentration. Lemna aequinoctialis had a high growth rate and strong phosphorus uptake capacity in low phosphorus water environment, suitable for low-phosphorus landscape water treatment.
(7) Hill equation had a higher precision compared with Michaelis-Menten equation, when correlating H2PO4-uptake rate of three duckweed species and phosphorus concentration in solution. When simulating the relation curve between H2PO4-uptake rate of Spirodela polyrrhiza and phosphorus concentration in solution under different environmental conditions, Hill equation showed a better result than Michaelis-Menten equation. The relationship between H2PO4-uptake rate of duckweed and phosphorus concentration in solution followed a sigmoid curve.
(1)稀脉浮萍的相对生长率RGR高于少根紫萍和紫萍,稀脉浮萍叶状体增殖速度最快。在实验室模拟培养条件下,稀脉浮萍叶状体的倍增时间最短,为3.23,少根紫萍和紫萍叶状体倍增时间分别为4.15天和4.81天。单位鲜重稀脉浮萍吸收的磷量显著低于少根紫萍和紫萍,其高磷累积吸收量建立在生物量大量扩增基础上。稀脉浮萍对H2PO4-的的亲和力常数Km和最大吸收速率Vmax均最小,紫萍的Km和Vmax最大。稀脉浮萍对H2PO4-的亲和力更强,在较低的离子浓度下就可以达到Vmax；而当环境中磷浓度较高时,紫萍对磷的净化速率高于少根紫萍和稀脉浮萍。实验室模拟培养期间,紫萍体内的ACPase和H+-ATPase活力水平始终最高,稀脉浮萍次之,少根紫萍最低。从细胞学形态结构上分析,紫萍和少根紫萍叶绿体基粒内囊体片层结构排列紧密,叶绿体周围环绕多个线粒体,而稀脉浮萍叶绿体中存在较大的淀粉粒构和空泡结构,片层结构松散。紫萍和少根紫萍的细胞超微结构更有利于植物光能利用效率和同化物运输能力的提高。因此,紫萍对磷的吸收能力更强,更适于对高磷浓度污水进行修复。
(2)紫萍RGR随光照强度增加而逐渐上升,在60001x高光强下,紫萍叶状体仍维持较高扩增速度。紫萍累积吸收磷量随着光照强度提高而升高,但单位鲜重浮萍吸收的磷量在60001x高光强条件下最低,30001x光强时的磷吸收能力略高于15001x,但两者之间无显著差异。随着光照强度增加,紫萍对H2P04-的最大吸收速率Vmax增加；不同光强条件下紫萍对H2P04-的亲和力常数Km略有差异,3000lx和60001x光强下略高于15001x光强。随着光照强度的增加,紫萍体内ACPase活力增加。实验室模拟培养期间,紫萍H+-ATP酶活力呈先缓慢下降随后逐渐上升的现象。培养后期低光强下,紫萍的H+-ATP酶活力超过高光强处理。强光下,紫萍叶肉细胞内的叶绿体数明显增加；弱光时虽然叶绿体数较少,但其基粒片层结构变厚,增加了光合作用反应面积。
(3)25℃时紫萍RGR最高,35℃略低,10℃最低。低温对紫萍生长的胁迫作用甚于高温。紫萍的累积吸收磷量随着温度的升高而增加,但单位鲜重浮萍吸收的磷量在10℃时最高,35℃次之,25℃最低。25℃和35℃条件下,紫萍的高磷累积吸收量通过生物量的大量扩增实现。随着温度增加,紫萍对H2P04-的最大吸收速率Vmax增加：不同温度之间亲和力大小依次为：25℃>10℃>35℃。35℃条件下紫萍对H2PO4-的低亲和力可能是高温胁迫作用所致。随着温度增加,紫萍体内ACPase活力增加。但长时间的低温胁迫也会使紫萍体内ACP酶活力显著提高。不同温度条件下,紫萍体内H+-ATPase活力的大小依次为：25℃>10℃。从细胞超微结构来看,与25℃适温条件相比,10℃低温和35℃高温下,紫萍细胞均存在一定程度的逆境胁迫作用。表现为低温时部分叶绿体皱缩成带状,叶肉细胞之间出现较大空隙；高温时叶绿体外膜溶解,基质外渗,脂质小球数量增多。
(4)不同pH条件下,紫萍RGR大小依次为pH6>pH5>pH7>pH8>pH9。紫萍磷累积吸收量在pH6条件下最高,当pH>6时,随着pH升高,磷累积吸收量降低。pH对单位鲜重紫萍吸收的磷量无显著影响。不同pH条件下,紫萍对H2P04-的最大吸收速率Vmax大小依次为：pH6>pH7>pH8>pH5>pH9,亲和力常数Km大小依次为：pH5>pH9>pH8>pH7>pH6。不同pH条件培养较长时间后,紫萍H+-ATPase和ACPase活力在pH6、pH7和pH8条件下高于pH5和pH9。紫萍在pH5条件下叶肉细胞中的叶绿体数量明显减少,类囊体片层结构松散。在pH6和pH7条件下,紫萍叶绿体数量丰富,叶肉细胞中溶质和细胞器较多,类囊体片层排列整齐紧密,光合效率高。pH8和pH9条件下,紫萍植株受到高OH-伤害,尤其是在pH9时,叶肉细胞中的叶绿体数量急剧减少,叶绿体和类囊体膜崩解,形成大量脂质小球。
(5)紫萍在水体总磷浓度0.1mg/L～15mg/L范围内,能保持较高的RGR,累积吸收磷量和单位鲜重紫萍吸收磷量均随着磷浓度升高而增加,当水体中总磷浓度达到45mg/L以上时,紫萍生长受限,RGR较低,磷累积吸收量和单位鲜重紫萍吸收的磷量均显著下降。当水体中总磷浓度在0.3mg/L-3mg/L范围时,紫萍对水体中磷的去除效率最高(约70%)。当水体中总磷浓度低于0.3mg/L时,紫萍体内ACPase活力升高,但长时间的低磷条件使紫萍生长受抑,从而使低磷和高磷条件之间ACPase活力的差异减小。与高磷条件相比,低磷条件下紫萍H+-ATPase活力较低。当水体中磷浓度较低时,紫萍叶绿体中出现大量淀粉粒结构,光合产物运输受阻；提高水体环境中磷浓度,叶绿体中淀粉粒结构开始消失；当进一步提高磷浓度使水体中总磷浓度达到45mg/L时,紫萍细胞出现质壁分离现象,叶绿体结构受到破坏,表现为外膜溶解,类囊体片层排列紊乱,对营养物的同化效率降低。
(6)不同类型富营养化水体中,三种浮萍的磷吸收效率存在差异。稀脉浮萍在氮磷浓度较低的景观水体中,磷累积吸收量最高。由于磷累积吸收量与生物量几乎按比例同步增加,以鲜重为基础算得的单位鲜重浮萍吸收的磷量并没有增加。紫萍在氮磷浓度较高的农业废水和生活污水中,磷累积吸收量和单位鲜重浮萍吸收的磷量均高于少根紫萍和稀脉浮萍,表现出对高磷浓度水体的磷吸收优势。稀脉浮萍叶绿素含量在景观水体中最高,在生活污水中最低。少根紫萍和紫萍叶绿素含量始终随着水体中氮磷浓度上升而升高。三种浮萍蛋白质含量均随着水体中氮磷浓度的升高而升高。不同类型富营养化水体中,稀脉浮萍体内蛋白质含量始终高于少根紫萍和紫萍。紫萍体内ACPase和H+-ATPase活力随着水体中磷浓度的增加而升高,在磷浓度较高的农业废水和生活污水中紫萍的ACPase和H+-ATPase活力显著高于稀脉浮萍和少根紫萍。在高磷浓度水体中,紫萍对污水中磷的去除效果、生长速率以及对磷的吸收积累能力均高于低磷环境,适于修复高磷污染的生活污水和农业废水；稀脉浮萍在低磷浓度水体中的生长速率高、对磷的吸收能力强,适于低磷污染的景观水体修复。
(7)用Hill方程对三种品种浮萍吸收H2P04速率与溶液中磷浓度关系进行拟合,其拟合度都高于Michaelis-Menten方程。不同环境条件下(包括光照强度、温度、pH、水体中初始磷浓度水平),对比Hill方程和Michaelis-Menten方程拟合的紫萍吸收HPO42-的速率与溶液中总磷浓度的关系曲线发现,前者的拟合效果均不同程度上优于后者。说明在较宽的磷浓度范围内,浮萍对H2P04的吸收速率与溶液中磷浓度之间表现出“S形”吸收曲线关系。
With the fast development of economy and population, water body eutrophication has become one of the most important environmental problems. Phosphorus is known to play an important role in water eutrophication. In order to improve water quality it is crucial to remove phosphorus from eutrophic water effectively. In recent years, ecological remediation methods using aquatic plants have aroused widespread interest. One of the most efficient and economical ways is the use of duckweed for eutrophic water body treatment. However, studies to date mainly deal with duckweed growth capacity and mechanisms of nitrogen uptake, while knowledge concerning mechanisms of phosphorus uptake and controlling factors still remains limited. In this paper, phosphorus purification capacity of duckweed were studied by determining phosphorus uptake efficiency, uptake kinetics, characteristics of phosphorus metabolism enzymes in plants and cell ultrastructure changes under the influence of various factors(varieties, light intensity, temperature, pH, phosphorus concentration and water type). The absorption efficiency and mechanisms of phosphorus by duckweed under different conditions were determined, which provided an important theoretical reference for the development and utilization of duckweed in the phosphorus restoration of eutrophic water bodies. The main results were as follows:
(1) Lemna aequinoctialis had a higher relative growth rate than Spirodela oligorrhiza and Spirodela polyrrhiza, with a leaflets'doubling time of3.23days, while the doubling times of Spirodela oligorrhiza and Spirodela polyrrhiza were4.15and4.81days respectively. However, the phosphorus uptake capacity of Lemna aequinoctialis was siginificantly lower than that of Spirodela oligorrhiza and Spirodela polyrrhiza. The high phosphorus accumulation in Lemna aequinoctialis was a result of biomass increase. The Km and Vmax values for phosphorus uptake of Lemna aequinoctialis were lower than those of Spirodela oligorrhiza and Spirodela polyrrhiza, indicating a stronger affinity to phosphorus of Lemna aequinoctialis Spirodela polyrrhiza had the highest Km and Vmax values for phosphorus uptake, which might lead to a better performance of Spirodela polyrrhiza in purifying wastewater with a higher phosphorus concentration. Spirodela polyrrhiza had the highest ACPase and H+-ATPase activity, followed by Lemna aequinoctialis and Spirodela oligorrhiza. Morphologically, Spirodela oligorrhiza and Spirodela polyrrhiza had tightly arranged chloroplasts thylakoid membranes, and the chloroplasts were surrounded by many mitochondrias, while chloroplasts in Lemna aequinoctialis had relatively loose structure, with accumulated starch and vesicles. Cell ultrastructure of Spirodela oligorrhiza and Spirodela polyrrhiza was of great benefit to light use and transport of assimilation substance in plants. Therefore, Spirodela polyrrhiza had higher phosphorus absorption capability than Spirodela oligorrhiza and Lemna aequinoctialis, making it more suitable for purifying high-phosphorus polluted water.
(2) RGR value of Spirodela polyrrhiza increased with the increase of light intensity. Spirodela polyrrhiza fronds remained in a high expanding speed even in60001x high light intensity. Total absorbed phosphorus amount of Spirodela polyrrhiza increased with the increase of light intensity, while phosphorus uptake capacity was the lowest at60001x high light intensity. There was no significant difference between phosphorus uptake capacity with light intensity of30001x and15001x. With the increase of light intensity Vmax for phosphorus uptake of Spirodela polyrrhiza increased, while no change was observed for Km. Phosphorus affinity of Spirodela polyrrhiza with30001x and60001x light intensity were slightly higher than15001x. ACPase activity of Spirodela polyrrhiza also increased with the increase of light intensity. H+-ATPase activity firstly decreased, and then increased slowly during the cultivation period. At the later period of cultivation, H+-ATPase activity of Spirodela polyrrhiza with weak light intensity were higher than strong light intensity. When supplied with strong light intensity, chloroplasts in mesophyll cell of Spirodela polyrrhiza increased. Though the number of chloroplasts decreased with weak light intensity, grana lamella of chloroplasts thickened to increase its photosynthesis reaction area.
(3) The highest and lowest RGR values of Spirodela polyrrhiza were determined at25℃and10℃respectively. The growth of duckweed was negatively influenced more by low temperature than by high temperature. Total absorbed phosphorus in Spirodela polyrrhiza increased with temperature, while the highest phosphorus uptake capacity was detected at10℃, which indicated that biomass increase was the reason for high cumulation of absorbed phosphorus in25℃and35℃treatments. Vmax for phosphorus uptake of Spirodela polyrrhiza increased with the increase of temperature. Km value of phosphorus affinity changed at different temperatures:25℃>10℃>35℃. ACPase activity of Spirodela polyrrhiza increased with the increase of temperature, however, the stressing effect caused by a long period of time with low temperature may also lead to a higher phosphorus affinity. H+-ATPase activity of Spirodela polyrrhiza under different temperature was25℃>10℃>35℃. In terms of cell ultrastructure, Spirodela polyrrhiza was under stress to a certain degree in10℃and35℃treatments, as shown by shriveled. Chloroplasts and large cell cavity at10℃, and dissolved chloroplast envelope, exuded stroma and increased Plastoglobulus at35℃.
(4) RGR values of Spirodela polyrrhiza at different pH were pH6> pH5> pH7> pH8> pH9. The highest total cumulative absorbed phosphorus in Spirodela polyrrhiza was found at pH6, while phosphorus uptake capacity was not affected by pH. Vmax for phosphorus uptake of Spirodela polyrrhiza was pH6> pH7> pH8>pH5> pH9, and Km was pH5> pH9> pH8> pH7> pH6. H+-ATPase and ACPase activities of Spirodela polyrrhiza were higher at pH6, pH7and pH8than other treatments. The number of chloroplasts in mesophyll cell of Spirodela polyrrhiza decreased significantly at pH5, with loose strucutre of thylakoidas lamella. At pH6and pH7, Spirodela polyrrhiza had more chloroplasts, solvend and organelles, and tightly arranged thylakoids, which assured a high photosynthetic efficiency. Duckweed was damaged by high concentration of OH-under pH8and pH9treatments, especially at pH9. As a result, the number of chloroplasts decreased dramatically, while the membranes of chloroplasts and thylakoid were collapsed and many plastoglobulus ermerged.
(5) Spirodela polyrrhiza could maintain a high RGR in water with0.1mg/L to15mg/L of phosphorus. The growth of Spirodela polyrrhiza was restricted, and RGR value was low as phosphorus concentration in water was above45mg/L. The highest phosphorus removal efficiency of Spirodela polyrrhiza was reached approximately70%when phosphorus concentration in solution was ranged between0.3mg/L and3mg/L. ACPase activity of Spirodela polyrrhiza increased when phosphorus was lower than0.3mg/L. However, long time exposure to low phosphorus would limit the growth of Spirodela polyrrhiza, which thus reduced the difference of ACPase activities betweenlow and high phosphorus concentration treatments. H+-ATPase activity with high phosphorus treatment was higher than low phosphorus treatment. Under low phosphorus treatment, many starch grains appeared in chloroplasts and the transport of photosynthetic products was interrupted. As soon as phosphorus concentration of outer solution was increased, starch grains disappeared. When phosphorus concentration reached45mg/L, plasm olysis started and structure of chloroplasts was destroyed, indicated by disintegrated membranes, lamella of thylakoidas arranged in disorder, and decreased nutrients assimilatory efficiency.
(6) Difference existed in phosphorus uptake efficiency of three duckweek species in different types of eutrophic water. In landscape water with low nitrogen and phosphorus concentrations, total absorbed phosphorus in Lemna aequinoctialis was higher than in Spirodela oligorrhiza and Spirodela polyrrhiza. This is due to the increase of biomass. However, total absorbed phosphorus and phosphorus uptake capacity of Spirodela polyrrhiza was the highest in domestic sewage and agricultural waste water with higher nitrogen and phosphorus concentration. Spirodela polyrrhiza showed a high phosphorus uptake capacity in high phosphorus concentration environment. In landscape water, chlorophyll content of Lemna aequinoctialis was higher than the other two species, whereas in domestic sewage it was the lowest. With the increase of nitrogen and phosphorus concentration in water, chlorophyll content of Spirodela oligorrhiza and Spirodela polyrrhiza increasedand protein content also increasedin all three duckweed species. ACPase and H+-ATPase activity of Spirodela polyrrhiza, which increased with the increase of phosphorus concentration of outer water environment, was much higher than that of Lemna aequinoctialis and Spirodela oligorrhiza in domestic sewage and agricultural waste water treatments. In high phosphorus water environment, phosphorus removal efficiency, cumulative uptake capacity, and growth rate of Spirodela polyrrhiza were higher than low phosphorus environment, making it a better candidate for treating domestic sewage and agricultural waste water with a high phosphorus concentration. Lemna aequinoctialis had a high growth rate and strong phosphorus uptake capacity in low phosphorus water environment, suitable for low-phosphorus landscape water treatment.
(7) Hill equation had a higher precision compared with Michaelis-Menten equation, when correlating H2PO4-uptake rate of three duckweed species and phosphorus concentration in solution. When simulating the relation curve between H2PO4-uptake rate of Spirodela polyrrhiza and phosphorus concentration in solution under different environmental conditions, Hill equation showed a better result than Michaelis-Menten equation. The relationship between H2PO4-uptake rate of duckweed and phosphorus concentration in solution followed a sigmoid curve.
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