加拿大格兰德河水体磷素形态转化及水生生物对磷素吸收释放研究
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摘要
磷素是自然界重要的营养元素,在水生态系统中通常为首要的限制因子,其在水环境中的赋存、迁移和转化等过程对水生态系统的初级生产力具有重要意义,但过量的磷素进入水体后会严重影响水体质量,造成水体富营养化等一系列环境问题,因此充分了解水体磷素循环状况对于制定相关水体最佳养分管理措施具有十分重要的意义。目前对于水体磷素的研究逐渐从关注不同形态磷的浓度变化,转向定量描述磷素在不同形态间及在生物相-非生物相之间的转化过程,但对于某一具体河流在这方面的综合研究,例如水体可溶性有机磷(DOP)潜在生物可利用性及对水体活性磷库的贡献、精确定量正磷酸盐(P043-)被水生生物吸收和释放的过程,及占水体磷素绝大多数的悬浮颗粒态磷(PP)的潜在可利用活性的研究仍然较少,且还不完善,因此本研究从水体不同形态磷的生物可利用性角度,以汇入北美五大湖之一Eire湖的最大支流、位于加拿大安大略省的格兰德河流域为研究对象,采用同位素标记示踪技术和化学连续提取分级等方法,对不同磷形态季节性变化、相互转化关系、与水环境的关系及在被水生生物吸收释放等过程进行了研究,这将有助于加深对格兰德流域水体磷素循环过程的认识,加深了解磷素地球化学特性,为指导该流域生产的磷素养分管理,防止面源污染发生具有重要的理论意义和实践价值。主要结论如下：
1.格兰德河水体中可溶性有机磷参与磷素循环的研究
在格兰德中段、按水流方向设置三处采样点：布里奇波特(Bridgeport)、维多利亚(Victoria)和布莱尔(Blair)。主要探讨了水中可溶性有机磷(DOP)对水中可溶性生物活性磷(SRP)库的贡献、影响DOP浓度变化的特定酶种类及活性大小。这3个采样位点在本试验进行阶段水体DOP的平均浓度分别为11.24μg PL-1,16.73μgP L-1和40.19μg P L-1。在比较不同过滤压力条件(0mm Hg,100mm Hg,300mm Hg及500mm Hg)对水中DOP浓度影响时,发现水样的过滤压力对水体DOP浓度有显著影响(P<0.05),即过滤压力越大,滤液中的DOP浓度越高,这是由于在过滤过程中浮游动植物等水生生物部分断裂的肢体通过了滤膜进入滤液中成为DOP的一部分,或部分活性固体颗粒物的细胞膜在压力下破裂,使细胞液等含磷物质(主要为RNA等)渗入到滤液中从而影响了滤液中的DOP浓度；在探讨自然水体DOP对SRP库贡献的试验中,发现自然状态下水中DOP浓度随时间推移有逐渐减小的趋势,而水体SRP浓度有缓慢上升趋势,且不同采样季节格兰德河水体中的DOP的浓度下降速度与SRP浓度增加速率相似(都在小于2μg P L-1h-1范围变化)；在随后的水体酶活测定试验中发现格兰德河水体中存在的碱性磷酸酶在降解DOP再生为生物可利用活性磷的过程中发挥了及其重要的作用,表明DOP不但可以参与水体磷素的循环过程,而且是水体生物可利用活性磷的来源之一；同时也发现格兰德河水体胞外自由态碱性磷酸酶活性较强,占水体总碱性磷酸酶活性的50%左右；在DOP参与下的格兰德河水体磷素周转时间为10-24h,对比之前学者们的研究结果可知格兰德河是水体营养循环速度很快,水体生物具有较强的光合作用和新陈代谢活动旺盛的一条富营养化河流。
2.格兰德河水体中不同磷形态的季节变化及其与水环境的关系
格兰德河干流与支流水体总磷(TP)在2012年的平均浓度分别为87.42和82.15μg P L-1,单从水体TP浓度来说这两条河流都属于富营养化河流。相关性显示康内斯托加河水体TP浓度变化对于格兰德水体TP变化间没有显著影响(P<0.05),表明支流水体虽然携带较高磷浓度汇入干流,并不对水体磷素营养浓度产生决定性的影响,表明格兰德河中段水体磷素除了来源于支流输入以外,还有其它含较高磷素浓度的输入源,可能与支流河口位置与格兰德河中段采样位点间存在直接向格兰德河排放含有大量营养元素污水的滑铁卢市污水处理厂有关；格兰德河中段水体与支流康内斯托加河口处SRP和DOP全年平均浓度分别为9.77和10.14μg PL-1,10.85μg P L-1和16.28μg P L-1,相关性分析显示支流水体SRP和DOP浓度对于流水体相应指标无显著影响(P>0.05),表明格兰德河中段水体的溶解态磷(SRP和DOP)除来源于支流输入以外,在支流河口与格兰德河中段采样位点之间的河段还存在大量外源性溶解态磷输入；格兰德河干流与支流水体悬浮颗粒态磷(PP)全年平均浓度分别为67.51和55.30μg P L-1,相关性分析显示格兰德河支流河口水体颗粒态磷浓度对干流水体颗粒态磷浓度存在显著影响(P<0.05),可推测占格兰德河支流水体总磷全年平均百分比69.09%的悬浮颗粒态磷在汇入干流后可以随水流向下游移动,对格兰德河中段干流水体悬浮颗粒态磷贡献明显,格兰德河中段区域水体颗粒态磷主要来源于支流的输入。
相关性分析显示格兰德河水体PP与DOP浓度之间存在显著相关关系(P<0.05),表明藻类等水生有机体是兰德河水体PP重要组分,当藻类等浮游生物生长旺盛时,部分小型水生生物或其肢体在过滤水样时可以通过滤膜进入滤液成为DOP的一部分,而部分大型藻类等水生生物会被滤膜拦截成为悬浮颗粒态含磷物质的一部分;PP与SRP之间也存在极显著相关关系,表明PP可以作为潜在活性磷成为水中生物可利用磷库SRP的重要补充。格兰德河水温对TP、PP、SRP和DOP均有显著影响,其中水温与TP呈现极显著负相关关系,表明水生生物在水温较高的条件下活性较强,对水体磷素的吸收利用需求增加,因此水中TP浓度会随着水温的增加而呈现下降趋势；由于细菌、藻类等水生生物活性会随着水中溶解氧含量的增加而增加,而本研究显示溶解氧与水体TP间呈现极显著相关关系(P<0.01),表明细菌、藻类等浮游水生有机体对格兰德河水体TP组成贡献较大。
3.格兰德河水体中悬浮颗粒态磷的潜在活性分析
用化学连续分级提取方法获得格兰德河干流与支流水体悬浮颗粒态磷5种可潜在活性磷组分年度平均浓度变化趋势都呈现：BD-SRP>85℃NaOH-SRP> NH4C1-SRP> HC1-SRP> NaOH-SRP;其中BD-SRP组分平均浓度最高,甚至比NaOH-SRP浓度高出几倍,由于BD-SRP代表了与铁离子、铝离子、锰离子等金属离子结合对水中氧化还原状态很敏感的磷素组分,因此推测该区域格兰德河干流和支流水体从陆地生态系统受纳的含磷颗粒态物质的地球化学性质相似,它们流域的土壤可能都是由含铁、锰、镁和铝等金属离子相对丰富的基性岩母质发育而来；相关性分析显示,格兰德河水体悬浮颗粒态磷各可提取组分中,NH4CI-SRP浓度与水体总悬浮颗粒态磷浓度间存在良好的相关关系,表明代表不稳定结合态生物活性磷的NH4CI-SRP可以随着水体悬浮PP的增加,对水体生物可利用活性磷库产生一定影响；同时85℃NaOH-SRP与其它可提取组分相比浓度相对较高,且与水体DOP之间存在良好地相关关系,也说明格兰德河水体悬浮颗粒态磷大多以有机态形式存在,间接表明格兰德河水中细菌、藻类、原生动物等水生生物由于接受充足的光照和能量而生长旺盛、数量较多、活性较强、生物量丰富；格兰德河干流与支流水体悬浮颗粒态磷5种潜在活性组分中浓度最低的潜在活性磷组分都为NaOH-SRP,由于此种磷素组分代表了存在于腐殖质中或富集在微生物体内的含磷有机物,因此可以推断格兰德河干流与支流水体中与土壤、岩石结合的小型细菌藻类等有机颗粒物、水中腐殖质含量较少;格兰德河干流与支流水体悬浮颗粒态磷各潜在活性磷组分平均浓度高低值出现在不同季节,表明格兰德河干流域支流水环境条件存在一定差异,因为在不同水环境条件下水体腐殖质含量、细菌藻类等水生生物种群及数量会有所不同,水体悬浮颗粒态磷中有机组分含量随季节变化会存在差异,进而对水体悬浮颗粒态磷中潜在可利用磷素组分的释放过程和释放量产生影响。
4.水生生物对格兰德河水体中磷素的吸收和释放过程研究
采用同位素示踪技术(32P)探讨了河流生态系统中有关正磷酸盐循环的几个方面内容：格兰德河水体两种不同形态水生生物：河床石面水生生物(包括生长在河底卵石表面的表层藻类及微型底栖动物等)、悬浮颗粒态水生生物(细菌蓝藻等浮游生物)对水中被32P标记的正磷酸盐(32P-P043-)的吸收状态的研究结果显示在加入32P-PO43-后的前两个小时内河床石面生物和悬浮颗粒态水生生物体对32P的吸收速度很快,但随着时间推移河床石面生物逐渐占水中32P-PO43-吸收量的绝大多数,水中悬浮颗粒态水生生物体内所含32P的量由于被河床石面水生生物等其他水生生物吞食等原因呈下降趋势。水中河床石面生物和悬浮颗粒态水生生物在温度较高的夏季8月份对水中正磷酸盐(PO43-)的吸收速率最大,分比为0.33和0.24μg Pcm-2h-1；冬季12月份水中河床石面生物和悬浮颗粒态水生生物对水体正磷酸盐(PO43-)的吸收速率最小,分别为0.036和0.034μg P cm-2h-1。在对水体中存在的不同尺寸水生生物对PO43-的吸收贡献大小的研究发现无论季节如何变化,格兰德河水中尺寸在0.2～2μm的超微浮游生物(细菌)都是格兰德河水体吸收正磷酸盐(PO43-)的主要水生生物种类。水中正磷酸盐(PO43-)的周转时间呈现温度较高周转时间越短趋势,在夏季8月份最短,大概在0.75h左右,冬季相对较长大概在2.67h左右。用凝胶柱层析法对水体32P-PO43-被水生生物吸收利用后释放再生的形态的研究结果表明格兰德河水生生物释放32P磷的形态主要以小分子量(≥200MW)溶解态PO43-为主。
综上所述,本文对格兰德河干流与支流水生态系统各种形态磷素在不同季节的变化及不同磷形态之间之间相互转化情况,对目前研究较少的水体可溶性活性磷和水中悬浮颗粒态磷的潜在生物可利用性进行了探讨,并定量描述了格兰德河水体正磷酸盐在生物相和非生物相之间的转化情况,为制定该流域磷素养分最佳管理措施提供了科学依据。
Compared with other nutrient elements which are essential for life, such as carbon, hydrogen and sulphur etc. under natural conditions, phosphorus is always the main limiting element for aquatic ecosystems. Such changes as its emission, migration and transformation are of great significance to the primary productivity of aquatic ecosystems. In this study, the Grand River, Ontario, Canada, the largest tributary of one of the Great Lakes, Eire, was chosen to evaluate the relationship between the seasonal variation of different forms of phosphorus and water physicochemical properties. Such methods as the isotope labeling and chemical sequential extraction and so on were applied to explore the activity of the three specific forms of phosphorus in water, that is, the dissolved organic phosphorus (DOP), orthophosphate (PO43-) and suspended particulate phosphorus (PP), and their involvement in water phosphorus cycling for better understanding of the three major processes of the water phosphorus cycling in watershed of the Grand River:the absorption and release of PO43-from of the plankton and the benthos, the acquisition of the conceptual model based on the involvement of the phosphorus in such aquatic organisms as the phytoplankton and bacteria by being prey to the zooplankton and the protozoa, as well as the contribution of the potentially biological available phosphorus released from the DOP and suspended PP due to changes in environmental conditions to the total phosphorus in water.
The study is mainly composed of four parts.
The first part is about the activity of the DOP in rivers and its involvement in water phosphorus cycling. Three sampling points were chosen in the midsection of the Grand (?) from the upstream to the downstream:Bridgeport, Victoria and Blair, to investigate the contribution of the DOP in water to the soluable reactive phosphorus (SRP), and the specific species of enzyme influencing the DOP concentration in water and their activities. The average DOP concentration in the three sampling points was11.24μg P L-1,16.73μg P L-1and40.19μg P L-1respectively. Due to the different exogenous input such as the sewage discharge and disposal from the sewage treatment plants etc., these three sampling points represent the natural water environment under different nutrition conditions. Comparing the influence of different filtration pressures (0mm Hg,100mm Hg,300mm Hg and500mm Hg) on the DOP concentration in water, it was found that there was a positive correlation between the filtration pressures of water samples and the DOP concentration of the filtrates (P<0.05), that is, the larger the filtration pressure, the higher the concentration of the DOP in the filtrates, which may due to the fact that some of the limbs of the zooplankton and phytoplankton, immersed into the filtrates through the filter membrance and became part of the DOP and also the cell membrance of part of active solid particles bursted under pressure, resulting in the penetration of such phosphorus substances as cell sap (mainly for RNA etc.) into the filtrates and influencing the DOP concentration in the filtrates. Exlporing the contribution of the DOP in natural water to the SRP, it was shown that the DOP concentration in natural water decreased gradually over time,and besides the decline of the DOP concentration in water of the Grand River in different sampling seasons was similar with the increase of the SRP (They changed similarly at the range less thatn2ug PL-1h-1). It was also shown through the following water enzyme assay that the alkaline phosphatase (AP) in water of the Grand River played an important part in the DOP degradation into the PO43-which indicates that the DOP can participate in the water phosphorus cycling and futhermore it is one source of the water SRP. Besides, the AP existed not only in bodies of the plankton in water but also in the extracellular free forms with strong activity in water, acconting for about50%of the total AP activity of the water body. With the involvement of the DOP, the turnaround time of water phosphorus in water of the Grand River was10to24hours. Thus it could be got through the comparison of the experimental results from former studies that the Grand River is one river of eutrophication, with fast water nutrient cycling, strong photosynthesis and active metabolism for the aquatic life.
The second part discusses the relationship between the seasonal variation of different forms of phosphorus in water and water physicochemical properties. The watershed in the midsection of the Grand River and the Conestogo River, one important tributary of the Grand River, was studied. Results indicated that the average concentration of the total phosphorus (TP) in water of the Grand River and the Conestogo River in2012was individually87.42μg P L-1and82.15μg PL-1, which indicates clearly that these two rivers are of eutrophication. The correlation analysis indicated that the TP concentration variation in water of the Conestogo River had no impact on the TP in the Grand River (p=0.061), which implies that although the phosphorus in the water of the Conestogo River flowed into the main stream in the midsection it did not bring great impact on the nutrient concentration of water phosphorus. Thus it can be inferred that besides the input from the tributaries, for the water phosphorus in the midsection of the Grand River there are other input sources with high phosphours concentration, which may be due to the fact that there are sewage treatment plants of Waterloo which discharge waste water with abuandant nutrients directly into the Grand River at about15km along the river between the river mouth of the Conestogo River and the Bridgeport, the midsection of the Grand River. The annual average concentration of the SRP and the DOP in water of the midsection of the Grand River was9.77μg P L-1and10.85μg P L-1. That in the river mouth of the Conestogo River was10.14μg P L-1and16.28μg P L-1respectively. The correlation analysis showed that there was no good correlation between the SRP concentration and the DOP concentration at these two sampling points (P>0.05), which infers that besides the input from the Conestogo River, for the soluable phosphorus (SRP and DOP) there are input of the exogenous soluable phosphorus into water of the Grand River. The annual average concentration of the PP in water of the Grand River and the Conestogo River was67.51μg P L-1and55.30μg P L-1. It can be found through the correlation analysis that there was good correlation between the concentration of the suspended PP in midsection of the Grand River and the suspended PP concentration in the river mouth of the Conestogo River (P<0.05), thus it can be inferred that the suspended PP, accounting for67.09%of the TP in water of the Conestogo River, entered into the water body of the Grand River, flowed into the downstream and did great contribution to the suspended PP in the Grand River. The suspended PP in watershed of the midsection of the Grand River mainly came from the input from the tributaries.
The third part is the geochemical properties analysis of suspended particulate phosphorus in water.Chemical approaches could be used to extract5kinds of potentially mobile phosphorus in the water particulate phosphorus in the Grand River and its tributary, Conestogo River. The variation trend of these five kinds was BD-SRP>85℃NaOH-SRP> NH4C1-SRP> HC1-SRP> NaOH-SRP,which indicates that the average concentration of BD-SRP in suspended particulate phosphorus was the highest, and it was even several times higher than that of NaOH-SRP. As the BD-SRP was the phosphorus fragment which combined with such metal ions as the iron ion, the aluminum ion and the manganese ion etc., and was quite sensitive to the redox state in water, it could be speculated that in this area in water of the main stream of the Grand River as well as its tributaries, the geochemical properties of the substances with phosphorus particles received from the terrestrial ecosystems were similar. It was possible that the soil in the watershed developed from the basic rock parent material abudant in such metal ions as iron, magnesium manganese and aluminum and so on. The correlation analysis showed that in each extraction fragment from the suspended particulate phosphorus in water of the Grand River, there was a good correlation between the NH4C1-SRP and the total water suspended PP, implying that with the increase of the water suspended PP, the NH4C1-SRP, representing bioactive phosphorus in instable combination, could make the potentially mobile phosphorus in water increase accordingly. Simultaneously, compared with the concentration of other extraction fragments, that of the85℃NaO-SRP was relatively high and it was in good correlation with the DOP in water, which stated that the water suspended PP in the Grand River mostly existed in organic forms and indicated that in Grand River the biomass was abundantly available and the activity of the aquatic life was strong. In the five potentially active fragments the concentration of the NaOH-SRP was the lowest. As it represented phosphorus organisms existing in humus or gathering in microorganisms, it could be concluded that water of the Grand River and the Conestogo River, was poor in such organic particulate matters as small bacteria and algae combined with soil and rock,humus and small bacteria and algae absorbed by the humus. The activity of the aquatic life in particles such as the algae etc. in water of the Grand River and the Conestogo River was strong. Such aquatic life as the bacteria, the alge and the protozoa were abundant and grew vigorously. As in different water bodies, the water environment such as the population quantity of the aquatic life (for example, the conten of the humus, the bacteria and the algae etc.) may have impact on the release and the absorption of the potentially available phosphorus fragments in water suspended PP with the seasonal variation, the high and low value of the average concentration of each potentially active phosphorus fragment in the water suspended PP in Grand River and the Conestogo River were various in different seasons. Thus it could be conculded that for the Grand River there was difference in the water environment between the main stream watershed and the watershed of its tributaries
The fourth part concerns the absorption and release of orthophosphate by the aquatic life. The isotope labeling (32P) was used to discuss several aspects of the orthophosphate cycling in river ecosystem. In water of the Grand River, there are two different kinds of aquatic lives:the aquatic organisms at the surface of pebbles in the riverbed (including the suface algae living at the surface of pebbles at the bottom of rivers as well as the microbenthons), and the aquatic lives in suspended particles (such plankton as the blue algae). It was showed through the experiment about the absorption of the orthophosphate labelled by the32P,(namely32P-PO43-) in water that the amount of the32P in these two kinds of aquatic lives increased sharply the first two hours after adding32P-PO43-.However, as time went on, the aquatic organisms at the surface of pebbles in the riverbed gradually accounted for the vast majority of the absorption of32P-PO43-in water and the amount of32P in the acquatic organisms in suspended particles in water decreased due to such factors as its absorption by the aquatic lives at the surface of pebbles in the riverbed and other aquatic organisms. The absorption rate of orthophosphate (PO43-) for the epilithion of the riverbed and acquatic organisms in suspended paritcles in water was the maximum in such season as summer with high to(?)perature,especially in August, namely0.33and0.24μg P cm-2h-1and in December it was quite the opposite with the absorption rate as0.036and0.034μg P cm-2h-1During the study of the contribution of aquatic lives with different sizes in water to the absorption of PO43-it could be found that despite the seasonal changes, the ultraplankton with the size from0.2to2μm in water of the Grand River was the main acquatic species absorbing the PO43-. Furthermore, the trunaround time of PO43-in water decreased with the increase of temperature. In August, it was the shortest as0.75h and in winter it was relatively longer as2.67h. Gel chromatography was used to explore the release and regeneration form of the32P-PO43-in water abosrbed and utilized by the aquatic lives. It could be found that the release of32P by the acquatic lives in the Grand River was mainly in the form of the dissolved PO43-with small molecular weight (≥200MW).
To sum up, the study concerns the seasonal variation of different forms of phsophorus in the water ecosystem, their transformation organisms, the potential bioavailability of the suspended PP in water and the transformation between biofacies and non-biofacies of the orthophosphate and so on. The conceptual model of water phosphorus cycling was put forward to better understand the water phosphorus cycling in the Grand River, help with the ecological environment protection in the watershed of the Grand River and provide reference for similar studies about water phosphorus cycling in rivers. However, it must be mentioned that water in rivers is in a constant state of motion and variation. Thus there are massive changes for the water ecology. Different conclusions can be drawn at different time, so only by continuous and long-term monitoring and study in the river ecosystem can rules and conclusions be drawn.
1.格兰德河水体中可溶性有机磷参与磷素循环的研究
在格兰德中段、按水流方向设置三处采样点：布里奇波特(Bridgeport)、维多利亚(Victoria)和布莱尔(Blair)。主要探讨了水中可溶性有机磷(DOP)对水中可溶性生物活性磷(SRP)库的贡献、影响DOP浓度变化的特定酶种类及活性大小。这3个采样位点在本试验进行阶段水体DOP的平均浓度分别为11.24μg PL-1,16.73μgP L-1和40.19μg P L-1。在比较不同过滤压力条件(0mm Hg,100mm Hg,300mm Hg及500mm Hg)对水中DOP浓度影响时,发现水样的过滤压力对水体DOP浓度有显著影响(P<0.05),即过滤压力越大,滤液中的DOP浓度越高,这是由于在过滤过程中浮游动植物等水生生物部分断裂的肢体通过了滤膜进入滤液中成为DOP的一部分,或部分活性固体颗粒物的细胞膜在压力下破裂,使细胞液等含磷物质(主要为RNA等)渗入到滤液中从而影响了滤液中的DOP浓度；在探讨自然水体DOP对SRP库贡献的试验中,发现自然状态下水中DOP浓度随时间推移有逐渐减小的趋势,而水体SRP浓度有缓慢上升趋势,且不同采样季节格兰德河水体中的DOP的浓度下降速度与SRP浓度增加速率相似(都在小于2μg P L-1h-1范围变化)；在随后的水体酶活测定试验中发现格兰德河水体中存在的碱性磷酸酶在降解DOP再生为生物可利用活性磷的过程中发挥了及其重要的作用,表明DOP不但可以参与水体磷素的循环过程,而且是水体生物可利用活性磷的来源之一；同时也发现格兰德河水体胞外自由态碱性磷酸酶活性较强,占水体总碱性磷酸酶活性的50%左右；在DOP参与下的格兰德河水体磷素周转时间为10-24h,对比之前学者们的研究结果可知格兰德河是水体营养循环速度很快,水体生物具有较强的光合作用和新陈代谢活动旺盛的一条富营养化河流。
2.格兰德河水体中不同磷形态的季节变化及其与水环境的关系
格兰德河干流与支流水体总磷(TP)在2012年的平均浓度分别为87.42和82.15μg P L-1,单从水体TP浓度来说这两条河流都属于富营养化河流。相关性显示康内斯托加河水体TP浓度变化对于格兰德水体TP变化间没有显著影响(P<0.05),表明支流水体虽然携带较高磷浓度汇入干流,并不对水体磷素营养浓度产生决定性的影响,表明格兰德河中段水体磷素除了来源于支流输入以外,还有其它含较高磷素浓度的输入源,可能与支流河口位置与格兰德河中段采样位点间存在直接向格兰德河排放含有大量营养元素污水的滑铁卢市污水处理厂有关；格兰德河中段水体与支流康内斯托加河口处SRP和DOP全年平均浓度分别为9.77和10.14μg PL-1,10.85μg P L-1和16.28μg P L-1,相关性分析显示支流水体SRP和DOP浓度对于流水体相应指标无显著影响(P>0.05),表明格兰德河中段水体的溶解态磷(SRP和DOP)除来源于支流输入以外,在支流河口与格兰德河中段采样位点之间的河段还存在大量外源性溶解态磷输入；格兰德河干流与支流水体悬浮颗粒态磷(PP)全年平均浓度分别为67.51和55.30μg P L-1,相关性分析显示格兰德河支流河口水体颗粒态磷浓度对干流水体颗粒态磷浓度存在显著影响(P<0.05),可推测占格兰德河支流水体总磷全年平均百分比69.09%的悬浮颗粒态磷在汇入干流后可以随水流向下游移动,对格兰德河中段干流水体悬浮颗粒态磷贡献明显,格兰德河中段区域水体颗粒态磷主要来源于支流的输入。
相关性分析显示格兰德河水体PP与DOP浓度之间存在显著相关关系(P<0.05),表明藻类等水生有机体是兰德河水体PP重要组分,当藻类等浮游生物生长旺盛时,部分小型水生生物或其肢体在过滤水样时可以通过滤膜进入滤液成为DOP的一部分,而部分大型藻类等水生生物会被滤膜拦截成为悬浮颗粒态含磷物质的一部分;PP与SRP之间也存在极显著相关关系,表明PP可以作为潜在活性磷成为水中生物可利用磷库SRP的重要补充。格兰德河水温对TP、PP、SRP和DOP均有显著影响,其中水温与TP呈现极显著负相关关系,表明水生生物在水温较高的条件下活性较强,对水体磷素的吸收利用需求增加,因此水中TP浓度会随着水温的增加而呈现下降趋势；由于细菌、藻类等水生生物活性会随着水中溶解氧含量的增加而增加,而本研究显示溶解氧与水体TP间呈现极显著相关关系(P<0.01),表明细菌、藻类等浮游水生有机体对格兰德河水体TP组成贡献较大。
3.格兰德河水体中悬浮颗粒态磷的潜在活性分析
用化学连续分级提取方法获得格兰德河干流与支流水体悬浮颗粒态磷5种可潜在活性磷组分年度平均浓度变化趋势都呈现：BD-SRP>85℃NaOH-SRP> NH4C1-SRP> HC1-SRP> NaOH-SRP;其中BD-SRP组分平均浓度最高,甚至比NaOH-SRP浓度高出几倍,由于BD-SRP代表了与铁离子、铝离子、锰离子等金属离子结合对水中氧化还原状态很敏感的磷素组分,因此推测该区域格兰德河干流和支流水体从陆地生态系统受纳的含磷颗粒态物质的地球化学性质相似,它们流域的土壤可能都是由含铁、锰、镁和铝等金属离子相对丰富的基性岩母质发育而来；相关性分析显示,格兰德河水体悬浮颗粒态磷各可提取组分中,NH4CI-SRP浓度与水体总悬浮颗粒态磷浓度间存在良好的相关关系,表明代表不稳定结合态生物活性磷的NH4CI-SRP可以随着水体悬浮PP的增加,对水体生物可利用活性磷库产生一定影响；同时85℃NaOH-SRP与其它可提取组分相比浓度相对较高,且与水体DOP之间存在良好地相关关系,也说明格兰德河水体悬浮颗粒态磷大多以有机态形式存在,间接表明格兰德河水中细菌、藻类、原生动物等水生生物由于接受充足的光照和能量而生长旺盛、数量较多、活性较强、生物量丰富；格兰德河干流与支流水体悬浮颗粒态磷5种潜在活性组分中浓度最低的潜在活性磷组分都为NaOH-SRP,由于此种磷素组分代表了存在于腐殖质中或富集在微生物体内的含磷有机物,因此可以推断格兰德河干流与支流水体中与土壤、岩石结合的小型细菌藻类等有机颗粒物、水中腐殖质含量较少;格兰德河干流与支流水体悬浮颗粒态磷各潜在活性磷组分平均浓度高低值出现在不同季节,表明格兰德河干流域支流水环境条件存在一定差异,因为在不同水环境条件下水体腐殖质含量、细菌藻类等水生生物种群及数量会有所不同,水体悬浮颗粒态磷中有机组分含量随季节变化会存在差异,进而对水体悬浮颗粒态磷中潜在可利用磷素组分的释放过程和释放量产生影响。
4.水生生物对格兰德河水体中磷素的吸收和释放过程研究
采用同位素示踪技术(32P)探讨了河流生态系统中有关正磷酸盐循环的几个方面内容：格兰德河水体两种不同形态水生生物：河床石面水生生物(包括生长在河底卵石表面的表层藻类及微型底栖动物等)、悬浮颗粒态水生生物(细菌蓝藻等浮游生物)对水中被32P标记的正磷酸盐(32P-P043-)的吸收状态的研究结果显示在加入32P-PO43-后的前两个小时内河床石面生物和悬浮颗粒态水生生物体对32P的吸收速度很快,但随着时间推移河床石面生物逐渐占水中32P-PO43-吸收量的绝大多数,水中悬浮颗粒态水生生物体内所含32P的量由于被河床石面水生生物等其他水生生物吞食等原因呈下降趋势。水中河床石面生物和悬浮颗粒态水生生物在温度较高的夏季8月份对水中正磷酸盐(PO43-)的吸收速率最大,分比为0.33和0.24μg Pcm-2h-1；冬季12月份水中河床石面生物和悬浮颗粒态水生生物对水体正磷酸盐(PO43-)的吸收速率最小,分别为0.036和0.034μg P cm-2h-1。在对水体中存在的不同尺寸水生生物对PO43-的吸收贡献大小的研究发现无论季节如何变化,格兰德河水中尺寸在0.2～2μm的超微浮游生物(细菌)都是格兰德河水体吸收正磷酸盐(PO43-)的主要水生生物种类。水中正磷酸盐(PO43-)的周转时间呈现温度较高周转时间越短趋势,在夏季8月份最短,大概在0.75h左右,冬季相对较长大概在2.67h左右。用凝胶柱层析法对水体32P-PO43-被水生生物吸收利用后释放再生的形态的研究结果表明格兰德河水生生物释放32P磷的形态主要以小分子量(≥200MW)溶解态PO43-为主。
综上所述,本文对格兰德河干流与支流水生态系统各种形态磷素在不同季节的变化及不同磷形态之间之间相互转化情况,对目前研究较少的水体可溶性活性磷和水中悬浮颗粒态磷的潜在生物可利用性进行了探讨,并定量描述了格兰德河水体正磷酸盐在生物相和非生物相之间的转化情况,为制定该流域磷素养分最佳管理措施提供了科学依据。
Compared with other nutrient elements which are essential for life, such as carbon, hydrogen and sulphur etc. under natural conditions, phosphorus is always the main limiting element for aquatic ecosystems. Such changes as its emission, migration and transformation are of great significance to the primary productivity of aquatic ecosystems. In this study, the Grand River, Ontario, Canada, the largest tributary of one of the Great Lakes, Eire, was chosen to evaluate the relationship between the seasonal variation of different forms of phosphorus and water physicochemical properties. Such methods as the isotope labeling and chemical sequential extraction and so on were applied to explore the activity of the three specific forms of phosphorus in water, that is, the dissolved organic phosphorus (DOP), orthophosphate (PO43-) and suspended particulate phosphorus (PP), and their involvement in water phosphorus cycling for better understanding of the three major processes of the water phosphorus cycling in watershed of the Grand River:the absorption and release of PO43-from of the plankton and the benthos, the acquisition of the conceptual model based on the involvement of the phosphorus in such aquatic organisms as the phytoplankton and bacteria by being prey to the zooplankton and the protozoa, as well as the contribution of the potentially biological available phosphorus released from the DOP and suspended PP due to changes in environmental conditions to the total phosphorus in water.
The study is mainly composed of four parts.
The first part is about the activity of the DOP in rivers and its involvement in water phosphorus cycling. Three sampling points were chosen in the midsection of the Grand (?) from the upstream to the downstream:Bridgeport, Victoria and Blair, to investigate the contribution of the DOP in water to the soluable reactive phosphorus (SRP), and the specific species of enzyme influencing the DOP concentration in water and their activities. The average DOP concentration in the three sampling points was11.24μg P L-1,16.73μg P L-1and40.19μg P L-1respectively. Due to the different exogenous input such as the sewage discharge and disposal from the sewage treatment plants etc., these three sampling points represent the natural water environment under different nutrition conditions. Comparing the influence of different filtration pressures (0mm Hg,100mm Hg,300mm Hg and500mm Hg) on the DOP concentration in water, it was found that there was a positive correlation between the filtration pressures of water samples and the DOP concentration of the filtrates (P<0.05), that is, the larger the filtration pressure, the higher the concentration of the DOP in the filtrates, which may due to the fact that some of the limbs of the zooplankton and phytoplankton, immersed into the filtrates through the filter membrance and became part of the DOP and also the cell membrance of part of active solid particles bursted under pressure, resulting in the penetration of such phosphorus substances as cell sap (mainly for RNA etc.) into the filtrates and influencing the DOP concentration in the filtrates. Exlporing the contribution of the DOP in natural water to the SRP, it was shown that the DOP concentration in natural water decreased gradually over time,and besides the decline of the DOP concentration in water of the Grand River in different sampling seasons was similar with the increase of the SRP (They changed similarly at the range less thatn2ug PL-1h-1). It was also shown through the following water enzyme assay that the alkaline phosphatase (AP) in water of the Grand River played an important part in the DOP degradation into the PO43-which indicates that the DOP can participate in the water phosphorus cycling and futhermore it is one source of the water SRP. Besides, the AP existed not only in bodies of the plankton in water but also in the extracellular free forms with strong activity in water, acconting for about50%of the total AP activity of the water body. With the involvement of the DOP, the turnaround time of water phosphorus in water of the Grand River was10to24hours. Thus it could be got through the comparison of the experimental results from former studies that the Grand River is one river of eutrophication, with fast water nutrient cycling, strong photosynthesis and active metabolism for the aquatic life.
The second part discusses the relationship between the seasonal variation of different forms of phosphorus in water and water physicochemical properties. The watershed in the midsection of the Grand River and the Conestogo River, one important tributary of the Grand River, was studied. Results indicated that the average concentration of the total phosphorus (TP) in water of the Grand River and the Conestogo River in2012was individually87.42μg P L-1and82.15μg PL-1, which indicates clearly that these two rivers are of eutrophication. The correlation analysis indicated that the TP concentration variation in water of the Conestogo River had no impact on the TP in the Grand River (p=0.061), which implies that although the phosphorus in the water of the Conestogo River flowed into the main stream in the midsection it did not bring great impact on the nutrient concentration of water phosphorus. Thus it can be inferred that besides the input from the tributaries, for the water phosphorus in the midsection of the Grand River there are other input sources with high phosphours concentration, which may be due to the fact that there are sewage treatment plants of Waterloo which discharge waste water with abuandant nutrients directly into the Grand River at about15km along the river between the river mouth of the Conestogo River and the Bridgeport, the midsection of the Grand River. The annual average concentration of the SRP and the DOP in water of the midsection of the Grand River was9.77μg P L-1and10.85μg P L-1. That in the river mouth of the Conestogo River was10.14μg P L-1and16.28μg P L-1respectively. The correlation analysis showed that there was no good correlation between the SRP concentration and the DOP concentration at these two sampling points (P>0.05), which infers that besides the input from the Conestogo River, for the soluable phosphorus (SRP and DOP) there are input of the exogenous soluable phosphorus into water of the Grand River. The annual average concentration of the PP in water of the Grand River and the Conestogo River was67.51μg P L-1and55.30μg P L-1. It can be found through the correlation analysis that there was good correlation between the concentration of the suspended PP in midsection of the Grand River and the suspended PP concentration in the river mouth of the Conestogo River (P<0.05), thus it can be inferred that the suspended PP, accounting for67.09%of the TP in water of the Conestogo River, entered into the water body of the Grand River, flowed into the downstream and did great contribution to the suspended PP in the Grand River. The suspended PP in watershed of the midsection of the Grand River mainly came from the input from the tributaries.
The third part is the geochemical properties analysis of suspended particulate phosphorus in water.Chemical approaches could be used to extract5kinds of potentially mobile phosphorus in the water particulate phosphorus in the Grand River and its tributary, Conestogo River. The variation trend of these five kinds was BD-SRP>85℃NaOH-SRP> NH4C1-SRP> HC1-SRP> NaOH-SRP,which indicates that the average concentration of BD-SRP in suspended particulate phosphorus was the highest, and it was even several times higher than that of NaOH-SRP. As the BD-SRP was the phosphorus fragment which combined with such metal ions as the iron ion, the aluminum ion and the manganese ion etc., and was quite sensitive to the redox state in water, it could be speculated that in this area in water of the main stream of the Grand River as well as its tributaries, the geochemical properties of the substances with phosphorus particles received from the terrestrial ecosystems were similar. It was possible that the soil in the watershed developed from the basic rock parent material abudant in such metal ions as iron, magnesium manganese and aluminum and so on. The correlation analysis showed that in each extraction fragment from the suspended particulate phosphorus in water of the Grand River, there was a good correlation between the NH4C1-SRP and the total water suspended PP, implying that with the increase of the water suspended PP, the NH4C1-SRP, representing bioactive phosphorus in instable combination, could make the potentially mobile phosphorus in water increase accordingly. Simultaneously, compared with the concentration of other extraction fragments, that of the85℃NaO-SRP was relatively high and it was in good correlation with the DOP in water, which stated that the water suspended PP in the Grand River mostly existed in organic forms and indicated that in Grand River the biomass was abundantly available and the activity of the aquatic life was strong. In the five potentially active fragments the concentration of the NaOH-SRP was the lowest. As it represented phosphorus organisms existing in humus or gathering in microorganisms, it could be concluded that water of the Grand River and the Conestogo River, was poor in such organic particulate matters as small bacteria and algae combined with soil and rock,humus and small bacteria and algae absorbed by the humus. The activity of the aquatic life in particles such as the algae etc. in water of the Grand River and the Conestogo River was strong. Such aquatic life as the bacteria, the alge and the protozoa were abundant and grew vigorously. As in different water bodies, the water environment such as the population quantity of the aquatic life (for example, the conten of the humus, the bacteria and the algae etc.) may have impact on the release and the absorption of the potentially available phosphorus fragments in water suspended PP with the seasonal variation, the high and low value of the average concentration of each potentially active phosphorus fragment in the water suspended PP in Grand River and the Conestogo River were various in different seasons. Thus it could be conculded that for the Grand River there was difference in the water environment between the main stream watershed and the watershed of its tributaries
The fourth part concerns the absorption and release of orthophosphate by the aquatic life. The isotope labeling (32P) was used to discuss several aspects of the orthophosphate cycling in river ecosystem. In water of the Grand River, there are two different kinds of aquatic lives:the aquatic organisms at the surface of pebbles in the riverbed (including the suface algae living at the surface of pebbles at the bottom of rivers as well as the microbenthons), and the aquatic lives in suspended particles (such plankton as the blue algae). It was showed through the experiment about the absorption of the orthophosphate labelled by the32P,(namely32P-PO43-) in water that the amount of the32P in these two kinds of aquatic lives increased sharply the first two hours after adding32P-PO43-.However, as time went on, the aquatic organisms at the surface of pebbles in the riverbed gradually accounted for the vast majority of the absorption of32P-PO43-in water and the amount of32P in the acquatic organisms in suspended particles in water decreased due to such factors as its absorption by the aquatic lives at the surface of pebbles in the riverbed and other aquatic organisms. The absorption rate of orthophosphate (PO43-) for the epilithion of the riverbed and acquatic organisms in suspended paritcles in water was the maximum in such season as summer with high to(?)perature,especially in August, namely0.33and0.24μg P cm-2h-1and in December it was quite the opposite with the absorption rate as0.036and0.034μg P cm-2h-1During the study of the contribution of aquatic lives with different sizes in water to the absorption of PO43-it could be found that despite the seasonal changes, the ultraplankton with the size from0.2to2μm in water of the Grand River was the main acquatic species absorbing the PO43-. Furthermore, the trunaround time of PO43-in water decreased with the increase of temperature. In August, it was the shortest as0.75h and in winter it was relatively longer as2.67h. Gel chromatography was used to explore the release and regeneration form of the32P-PO43-in water abosrbed and utilized by the aquatic lives. It could be found that the release of32P by the acquatic lives in the Grand River was mainly in the form of the dissolved PO43-with small molecular weight (≥200MW).
To sum up, the study concerns the seasonal variation of different forms of phsophorus in the water ecosystem, their transformation organisms, the potential bioavailability of the suspended PP in water and the transformation between biofacies and non-biofacies of the orthophosphate and so on. The conceptual model of water phosphorus cycling was put forward to better understand the water phosphorus cycling in the Grand River, help with the ecological environment protection in the watershed of the Grand River and provide reference for similar studies about water phosphorus cycling in rivers. However, it must be mentioned that water in rivers is in a constant state of motion and variation. Thus there are massive changes for the water ecology. Different conclusions can be drawn at different time, so only by continuous and long-term monitoring and study in the river ecosystem can rules and conclusions be drawn.
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