渤海浮游植物生态动力学模型研究
摘要
近二十年来生态系统动力学模型已在各个海域得到了广泛应用它们以不同时空分辨率量化海洋生态系中各过程的相互作用机制建立了包含不同复杂程度生物物理过程的生态模型本文首先综述了各种海洋生态模型的发展特点分析了生态系统动力学模型的一般结构总结了模型中的典型生物过程的函数表达以及环流/混合温度透明度太阳辐射等环境因子对海洋生态系的影响
利用历史资料分析了渤海浮游生态系基本特征分析了叶绿素初级生产力的季节变化和水平分布规律讨论了营养盐的分布特征分析了影响营养盐循环生物生长的物理过程变化特征在资料分析基础上建立一个基于氮磷营养盐循环并与一个成熟的三维斜压水动力模型耦合的三维渤海初级生产模型
分析讨论了模型中参数取值对模拟结果的影响讨论了参数敏感性和不同过程对海洋生态系的影响模型对于浮游植物最大生长率浮游植物基础呼吸率浮游植物死亡率浮游动物捕食率水底碎屑矿化率等参数变化比较敏感渤海浮游植物生产限制因子比较复杂生物生产处于光和氮磷营养盐的交替控制下若是光照条件和营养盐水平的变化不改变各限制性因子的相对限制关系时对系统的影响不大反之若是显著改变了限制关系则会导致系统比较大的波动这种情况下仅仅考虑一种营养盐是不适宜的模型当中至少应该同时包括氮磷营养盐循环过程浮游植物生物量年循环规律表现为生物量峰值出现时间和峰值的相对大小计算结果表明生物过程参数既能影响峰值出现时间又能影响峰值的大小而径流输入变化和水平对流物理过程只影响该峰值的相对大小而不改变峰值出现的时间可见系统中水温辐射垂直对流/混合及生物过程影响物质的局地变化而水平对流输运过程则将局地的变化向其邻近的水域传播从而影响物质的水平分布
在此基础上模拟了1982 年渤海营养盐叶绿素和初级生产力的季节变化和水平分布规律并且用1982 资料校验了模拟结果讨论了渤海碳和氮磷营养盐循环和收支规律
渤海叶绿素和营养盐季节变化反相营养盐是浮游植物生长的基础渤海营养盐经历了夏季的减少和秋季回升冬季积累浮游植物生物量变化对应于营养盐变化在春季随着温度升高不断增加到夏季出现浮游植物的高峰此后开始下降至12 月到达一年的最低值
渤海各个区域的初级生产力的升降趋势基本一致最高峰发生在夏季7-8月冬季12-1 月为一年的最低期莱州湾初级生产力呈双峰结构其余三个海湾为单峰结构温度对于渤海初级生产力春季峰值出现与否有很大影响初级生产力的水平分布不均匀四个海区中以莱州湾生产力最高中央海区和辽东湾次之渤海湾生产力远低于其他海域透明度低是造成渤海湾生产力低于其他海区的主要原因可见透明度和温度等物理因素的变化对渤海初级生产力的变化和分布有重要的作用
营养盐水平分布的季节变化特点是渤海中部变化相对比较平稳而其他三个湾变化波动则比较大尤其是在莱州湾冬季最高春秋水华后则最低磷酸盐冬
Great progress has been made in the research of ecosystem dynamic models during the last two decades. Physical-biological models of various levels of sophistication have been developed for different regions of the ocean. Firstly the features of different marine ecosystem models are summarized and discussed in this paper. Different functions of the representative biological process and the effect of circulation, water temperature, transparence and solar radiation on the ecosystem are described and analyzed.
Based on the historical data, this paper demonstrates the variation and distribution of chlorophyll-a, primary production and nutrient in the Bohai Sea and analyses the features of the physical process. According to the analysis of the historical data, a three-dimensional ecosystem model based on the cycles of phosphate and nitrate is developed for the Bohai Sea, which is coupled with a three-dimensional physical transport model.
Sensitivity analysis indicates that the variation of the phytoplankton biomass is sensitive to phytoplankton maximum growth rate, phytoplankton mortality rate, zooplankton grazing rate and remineralization rate of benthic detritus. Light, temperature and nutrient can all affect the production of phytoplankton. If their variations do not change the relation of the limitation, then the variations have little effect on the ecosystem; otherwise they can change the system greatly. The biological process and transparence can both affect the occurrence of the phytoplankton biomass peak and its amplitude. While the variation of the river discharge and the physical transport process can only change the relative amplitude of the peak, but have no effect on its occurrence. It can be concluded that transparence, solar radiation, biological process and the vertical mixing affect the local variation of the ecosystem, while the horizontal advection can spread the local variation of the system to adjacent field and thus affect the distribution of the phytoplankton and nutrient.
The model is then used to simulate the variation and distribution of chlorophyll-a, primary production and nutrient in 1982 and the simulation is validated by the data in 1982/1983. The concentration of both DIN and phosphate decreases from spring to summer and increases from autumn to winter in all the areas. The variation is a response to the consumption of the phytoplankton. In spring, the phytoplankton biomass increases as the temperature increasing and reaches the highest peak in summer. The concentration of nutrient drops to the lowest level during the same period. After the phytoplankton bloom of summer the dissolved inorganic nutrients increase gradually as the input of river increase and the decay of the thermocline.
The variations of the primary production are same in different regions of the Bohai Sea. The high value of primary production appears in July and August and the lowest value appears in December and January. The primary production in the
Laizhou Bay has two peaks and has one in the other part of the Bohai Sea in 1982. Water temperature has great influence on whether the peak of the primary production will appear in spring. Among the four parts of the Bohai Sea, the primary production is the highest in the Laizhou Bay and the lowest in Bohai Bay. The low transparence in the Bohai Bay is the main cause of the lowest primary production in it. So physical factors, such as temperature and transparence have a great effect on the variation and distribution of the primary production. The variation of the nutrient in the Central Bohai Sea is relatively stable while the variation is strong in coastal area of the Bohai Sea. The high concentration locates in the Bohai Bay in winter. The phosphate concentration in the northwest part of Liaodong Bay maintains a high level in the whole year. The value of phytoplankton biomass of the whole Bohai Sea is low in winter and the high value of the biomass first appears in the coastal area in spring. The high value expands from Laizhou Bay and Bohai Bay to the Central Bohai Sea in summer and the concentration of chlorophyll-a is about 1 mg/m3 in the Central Bohai Sea. The isoline of 1 mg/m3 moves to the coastal area again in autumn. The Huanghe river has great influence on the distribution of nitrate and phytoplankton concentration in the Laizhou Bay. The vertical distribution of nutrients is homogeneous in the whole year, except in the summer the concentration at the bottom layer is greater than that in the surface. This is due to the occurrence of thermocline in the Bohai Sea. The stratification does not decrease the concentration of chlorophyll-a in summer but has an effect on the vertical distribution of chlorophyll-a. The stability of water with enough supply of nutrient is propitious to the production of phytoplankton. Production and respiration are the most important sink and source of nutrients. The remineralization of the detritus pool is an important source of nutrient regeneration. It can compensate 23 percent of the consumption of nutrient from the production process. The net nutrient budget is -3.05 kilotons of P and 31.6 kilotons of N on the whole. The net carbon budget is 110 kilotons and the Bohai Sea is a weak sink of CO2 of the atmosphere. 13.7% of the gross primary production transfers to high trophic level. Remineralization at the bottom is the mechanism of transferring the nutrient from organic form to inorganic form. Finally a biological model, coupled with a three-dimensional physical transport model is described. Simulated distributions of plankton and nutrient are obtained from this model with nitrate input from a river. The simulations are then resampled and the data are used in numerical experiments to assess the ability of using an adjoint data assimilation approach for estimation the poorly known parameters of this ecosystem model. The spatial resolution and temporal resolution of the data are complementary in the assimilative model, thus the improvement of either of them can result in better recoveries of the model parameters. The assimilation of phytoplankton data is essential to recover the model parameters. Observational data collected during a Sino-German cooperation are taken as an example. Twin experiments, using simulated data of the same type and spatial and temporal distribution as that of the investigation, demonstrate the feasibility to estimating the model parameters with the
in situ observations. Some advice is given about the designing of sampling strategies for making measurements in a project like ours.
利用历史资料分析了渤海浮游生态系基本特征分析了叶绿素初级生产力的季节变化和水平分布规律讨论了营养盐的分布特征分析了影响营养盐循环生物生长的物理过程变化特征在资料分析基础上建立一个基于氮磷营养盐循环并与一个成熟的三维斜压水动力模型耦合的三维渤海初级生产模型
分析讨论了模型中参数取值对模拟结果的影响讨论了参数敏感性和不同过程对海洋生态系的影响模型对于浮游植物最大生长率浮游植物基础呼吸率浮游植物死亡率浮游动物捕食率水底碎屑矿化率等参数变化比较敏感渤海浮游植物生产限制因子比较复杂生物生产处于光和氮磷营养盐的交替控制下若是光照条件和营养盐水平的变化不改变各限制性因子的相对限制关系时对系统的影响不大反之若是显著改变了限制关系则会导致系统比较大的波动这种情况下仅仅考虑一种营养盐是不适宜的模型当中至少应该同时包括氮磷营养盐循环过程浮游植物生物量年循环规律表现为生物量峰值出现时间和峰值的相对大小计算结果表明生物过程参数既能影响峰值出现时间又能影响峰值的大小而径流输入变化和水平对流物理过程只影响该峰值的相对大小而不改变峰值出现的时间可见系统中水温辐射垂直对流/混合及生物过程影响物质的局地变化而水平对流输运过程则将局地的变化向其邻近的水域传播从而影响物质的水平分布
在此基础上模拟了1982 年渤海营养盐叶绿素和初级生产力的季节变化和水平分布规律并且用1982 资料校验了模拟结果讨论了渤海碳和氮磷营养盐循环和收支规律
渤海叶绿素和营养盐季节变化反相营养盐是浮游植物生长的基础渤海营养盐经历了夏季的减少和秋季回升冬季积累浮游植物生物量变化对应于营养盐变化在春季随着温度升高不断增加到夏季出现浮游植物的高峰此后开始下降至12 月到达一年的最低值
渤海各个区域的初级生产力的升降趋势基本一致最高峰发生在夏季7-8月冬季12-1 月为一年的最低期莱州湾初级生产力呈双峰结构其余三个海湾为单峰结构温度对于渤海初级生产力春季峰值出现与否有很大影响初级生产力的水平分布不均匀四个海区中以莱州湾生产力最高中央海区和辽东湾次之渤海湾生产力远低于其他海域透明度低是造成渤海湾生产力低于其他海区的主要原因可见透明度和温度等物理因素的变化对渤海初级生产力的变化和分布有重要的作用
营养盐水平分布的季节变化特点是渤海中部变化相对比较平稳而其他三个湾变化波动则比较大尤其是在莱州湾冬季最高春秋水华后则最低磷酸盐冬
Great progress has been made in the research of ecosystem dynamic models during the last two decades. Physical-biological models of various levels of sophistication have been developed for different regions of the ocean. Firstly the features of different marine ecosystem models are summarized and discussed in this paper. Different functions of the representative biological process and the effect of circulation, water temperature, transparence and solar radiation on the ecosystem are described and analyzed.
Based on the historical data, this paper demonstrates the variation and distribution of chlorophyll-a, primary production and nutrient in the Bohai Sea and analyses the features of the physical process. According to the analysis of the historical data, a three-dimensional ecosystem model based on the cycles of phosphate and nitrate is developed for the Bohai Sea, which is coupled with a three-dimensional physical transport model.
Sensitivity analysis indicates that the variation of the phytoplankton biomass is sensitive to phytoplankton maximum growth rate, phytoplankton mortality rate, zooplankton grazing rate and remineralization rate of benthic detritus. Light, temperature and nutrient can all affect the production of phytoplankton. If their variations do not change the relation of the limitation, then the variations have little effect on the ecosystem; otherwise they can change the system greatly. The biological process and transparence can both affect the occurrence of the phytoplankton biomass peak and its amplitude. While the variation of the river discharge and the physical transport process can only change the relative amplitude of the peak, but have no effect on its occurrence. It can be concluded that transparence, solar radiation, biological process and the vertical mixing affect the local variation of the ecosystem, while the horizontal advection can spread the local variation of the system to adjacent field and thus affect the distribution of the phytoplankton and nutrient.
The model is then used to simulate the variation and distribution of chlorophyll-a, primary production and nutrient in 1982 and the simulation is validated by the data in 1982/1983. The concentration of both DIN and phosphate decreases from spring to summer and increases from autumn to winter in all the areas. The variation is a response to the consumption of the phytoplankton. In spring, the phytoplankton biomass increases as the temperature increasing and reaches the highest peak in summer. The concentration of nutrient drops to the lowest level during the same period. After the phytoplankton bloom of summer the dissolved inorganic nutrients increase gradually as the input of river increase and the decay of the thermocline.
The variations of the primary production are same in different regions of the Bohai Sea. The high value of primary production appears in July and August and the lowest value appears in December and January. The primary production in the
Laizhou Bay has two peaks and has one in the other part of the Bohai Sea in 1982. Water temperature has great influence on whether the peak of the primary production will appear in spring. Among the four parts of the Bohai Sea, the primary production is the highest in the Laizhou Bay and the lowest in Bohai Bay. The low transparence in the Bohai Bay is the main cause of the lowest primary production in it. So physical factors, such as temperature and transparence have a great effect on the variation and distribution of the primary production. The variation of the nutrient in the Central Bohai Sea is relatively stable while the variation is strong in coastal area of the Bohai Sea. The high concentration locates in the Bohai Bay in winter. The phosphate concentration in the northwest part of Liaodong Bay maintains a high level in the whole year. The value of phytoplankton biomass of the whole Bohai Sea is low in winter and the high value of the biomass first appears in the coastal area in spring. The high value expands from Laizhou Bay and Bohai Bay to the Central Bohai Sea in summer and the concentration of chlorophyll-a is about 1 mg/m3 in the Central Bohai Sea. The isoline of 1 mg/m3 moves to the coastal area again in autumn. The Huanghe river has great influence on the distribution of nitrate and phytoplankton concentration in the Laizhou Bay. The vertical distribution of nutrients is homogeneous in the whole year, except in the summer the concentration at the bottom layer is greater than that in the surface. This is due to the occurrence of thermocline in the Bohai Sea. The stratification does not decrease the concentration of chlorophyll-a in summer but has an effect on the vertical distribution of chlorophyll-a. The stability of water with enough supply of nutrient is propitious to the production of phytoplankton. Production and respiration are the most important sink and source of nutrients. The remineralization of the detritus pool is an important source of nutrient regeneration. It can compensate 23 percent of the consumption of nutrient from the production process. The net nutrient budget is -3.05 kilotons of P and 31.6 kilotons of N on the whole. The net carbon budget is 110 kilotons and the Bohai Sea is a weak sink of CO2 of the atmosphere. 13.7% of the gross primary production transfers to high trophic level. Remineralization at the bottom is the mechanism of transferring the nutrient from organic form to inorganic form. Finally a biological model, coupled with a three-dimensional physical transport model is described. Simulated distributions of plankton and nutrient are obtained from this model with nitrate input from a river. The simulations are then resampled and the data are used in numerical experiments to assess the ability of using an adjoint data assimilation approach for estimation the poorly known parameters of this ecosystem model. The spatial resolution and temporal resolution of the data are complementary in the assimilative model, thus the improvement of either of them can result in better recoveries of the model parameters. The assimilation of phytoplankton data is essential to recover the model parameters. Observational data collected during a Sino-German cooperation are taken as an example. Twin experiments, using simulated data of the same type and spatial and temporal distribution as that of the investigation, demonstrate the feasibility to estimating the model parameters with the
in situ observations. Some advice is given about the designing of sampling strategies for making measurements in a project like ours.
引文
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