Research on the application of EwE model in aquatic ecosystems and a case study of Changtan Reservoir
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摘要:
EwE（Ecopath with Ecosim）模型是一种用于定量研究水生态系统食物网结构和能量流动特征的模型。总结了EwE模型中Ecopath、Ecosim、Ecospace和Ecotracer模块的原理，综述了该模型在不同类型水生态系统中的应用研究进展，发现Ecopath可用来评估海洋和淡水生态系统的成熟度并确定关键种和生态容量，明确浮游植物生产量（P）对生态系统总通量（TST）的重要贡献；Ecosim用于在时间尺度上揭示关键种捕捞、港口建设等人类活动对生态系统结构和功能的影响机制；Ecospace可用来阐明海上平台建设、发电厂运行、火山爆发等外部因素影响下渔业经济和生态系统结构的空间差异性；Ecostracer可用来追踪同位素、重金属和新型污染物等物质在食物网中的迁移过程。采用Ecopath分析了长潭水库的营养结构和能量流动，结果表明，长潭水库营养级介于1.000~3.093，食物网结构简单，TPP/TR为2.445，是一个相对成熟的生态系统，能量传递效率较低。未来应加强气候变化和人类活动对水生态系统结构演替的影响研究，推进新污染物在食物网中富集特征研究，为水域生态系统健康状态评估和渔业经济发展政策的调整提供支撑。
Abstract:Ecopath with Ecosim (EwE) model is commonly used to quantitatively study the food web structure and energy flow characteristics in the aquatic ecosystem. The principles and progress of Ecopath, Ecosim, Ecospace, and Ecotracer modules of EwE model and its application in different types of aquatic ecosystems were summarized. The results indicated that Ecopath could be used to estimate the maturity of the ocean and freshwater ecosystems, to determine key species and ecological capacity, and to clarify the important contribution of phytoplankton production (P) to total system throughput (TST) in the aquatic ecosystem. Ecosim was used to reveal the impact mechanisms of human activities (e.g. keystone species catching, port construction) on ecosystem structure and function on a time scale. Ecospace was used to clarify spatial variability of fishing activity and ecosystem structure under external factor effects (e.g. offshore platform construction, power plant operation, and volcanic eruption). Ecotracer was used to trace the immigration progress of isotopes, heavy metal, new pollutant, and other matters in the food web. Combined with Ecopath, the nutrient structure and energy flow of Changtan Reservoir were analyzed. The results showed that the trophic level was in the range of 1.000-3.093, indicating a simple food web structure. The total primary production/total respiration (TPP/TR) value was 2.445, indicating Changtan Reservoir was a relatively mature ecosystem, and it had a low energy transfer efficiency. In the future, studies on the impacts of climate change and human activities on the structural succession of aquatic ecosystems, and the enrichment characteristics of new pollutants in the food web should be strengthened to provide a scientific basis for the assessment of aquatic ecosystem health and the adjustment of fishery economic development policies.
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Key words:
- EwE model /
- aquatic ecosystems /
- climate change /
- new pollutants /
- Changtan Reservoir
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图 2 TST与生态系统中浮游植物 P 和浮游植物 B 的关系
Figure 2. Relationship of TST with the P and B of phytoplankton in ecosystem
图 3 表2应用实例中不同TE占比
Figure 3. Occupancy of different TE values across application examples in table 2
图 5 长潭水库林德曼能量流动图
注：P代表初级生产者，D代表碎屑，均为第Ⅰ营养级；Ⅰ、Ⅱ、Ⅲ、Ⅳ、Ⅴ为营养级。Q 为消费量； E 为净迁移量； Y 为捕捞量； B 为生物量；TST为系统总流量； R 为呼吸量；TE为能量传递效率；M2为捕食死亡率。
Figure 5. Lindeman spine of Changtan Resevoir’s energy flow
表 1 Ecopath主要参数及参数来源
Table 1. Main parameters and their resources of Ecopath
主要参数 参数含义 获取来源 取值范围 生物量（B） 各功能组在某个时间段
内的平均生物量来自实测值或者调查资料 无 生产量/生物量（P/B） 各功能组在某个时间段内的生产力 P/B为捕捞死亡率（F）和自然死亡率（M）之和，其中F根据实际捕捞量进行计算，M根据Pauly[14]的经验公式进行计算。鱼类P/B根据fishbase等网站计算得到 无 消费量/生物量（Q/B） 各功能组在某个时间段内的
消费能力采用实测值或参考临近水域或通过日消费比值进行估计或使用fishbase等网站估计。其中鱼类Q/B可通过Palomares等[15]的经验公式估计，无脊椎动物Q/B使用P/Q来代替 无 生产量/消费量（P/Q） 各功能组生产量和消费量的比值 Ecopath根据P/B和Q/B来计算或参考相关研究的取值 通常为0.05~0.30，根据
体重变化调整生态营养效率（EE） 各功能组生产量被利用的效率 Ecopath根据B、P/B和Q/B计算或根据相关研究估计 0~1，其取值
通常接近1饮食结构（DC） 各功能组营养级数值的确定 采用胃含物分析法和稳定同位素法测定。鱼类DC可通过fishbase等网站获取，浮游生物和底栖动物DC参考其他研究 各功能组DC比例之和为1 
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表 2 Ecopath在中国的应用实例
Table 2. Application examples of Ecopath in China
水体名称 模型应用年份 系统总流量(TST)/〔t/(km2·a)〕 TPP/TR 浮游植物的(P/B)/a−1 系统连接指数(CI) 太湖[21] 1991—1995 13 386 3.85 185 0.21 太湖[22] 2008—2009 66 245 4.22 410 0.19 太湖[23] 2017—2018 7 388 2.55 410 0.21 太湖竺山湾[24] 2015 10 145 2.37 185 0.24 五里湖[25] 2009 9 132 1.34 262 0.28 淀山湖[26] 2008—2009 4 099 2.80 185 0.19 巢湖[27] 2007—2010 41 003 13.53 185 0.20 千岛湖[28] 2000 24 271 1.99 201 0.23 千岛湖[29] 2016 24 698 6.51 180 0.26 南海北部湾[30] 1997—1999 11 006 3.18 231 0.33 杭州湾[31] 2006 18 958 2.67 476 0.31 长江口[32] 2004 6 342 2.53 200 0.54 长江口[33] 2016—2017 1 329 1.25 119 0.35 长江口及毗邻水域[34] 2000—2006 1 959~6 554 1.82~5.29 180~200 0.41~0.45 象山港[35] 2011—2014 2 227~2 229 1.52 180 0.34 渤海[36] 1982—1992 3 316~5 362 8.40~9.75 380~398 渤海[37] 2014—2015 10 499 5.38 250 0.33 俚岛人工礁区[38] 2009 10 787 1.84 71 0.20 獐子岛人工鱼礁区[39] 2010—2012 28 691~40 486 2.05~2.29 105~132 0.20~0.23 獐子岛海域[40] 2017—2018 17 007-17 738 1.79~2.05 140 0.22 海州湾[41] 2013 9 335 1.33 107 0.42 庙岛[42] 1998 3 172 2.47 100 0.44 枸杞岛海藻场[43] 2004—2008 28 019 1.25 119 0.33 三沙湾[44] 2012 2 344 2.77 105 0.40
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表 3 长潭水库Ecopath食物网模型的输入与输出参数
Table 3. Input and output parameters of Ecopath food web model of Changtan Reservoir
功能组 营养级 B/(t/km²) (P/B)/a−1 (Q/B)/a−1 EE (Q/P)/a−1 其他鱼类 2.547 9.021 1.200 13.000 0.500 0.092 鲌鱼 3.093 0.050 1.090 9.140 0.418 0.124 鲴鱼 2.000 0.351 2.011 16.270 0.950 0.124 鲤鱼 2.652 0.019 1.210 7.270 0.950 0.166 鲫鱼 2.247 5.931 1.150 8.370 0.950 0.137 鳙鱼 2.827 6.840 1.300 7.200 0.808 0.181 鲢鱼 2.248 9.290 1.500 8.450 0.800 0.178 草鱼 2.454 2.737 1.730 9.660 0.950 0.179 底栖动物 2.111 3.141 11.000 15.000 0.964 0.733 桡足类 2.170 5.270 6.000 100.000 0.935 0.060 枝角类 2.012 1.410 16.000 100.000 0.949 0.160 轮虫 2.050 3.350 40.000 150.000 0.950 0.267 大型植物 1.000 10.357 10.000 — 0.900 — 浮游植物 1.000 25.130 90.000 — 0.412 — 碎屑 1.000 50.000 — — 0.167 — 注：数据中加粗字体为模型输出值；—表示无须输入或输出，其余为输入值。
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