滆湖ecopath模型构建及围网放养鲢鳙的生态效应
摘要
滆湖是苏南地区第二大淡水湖，面积164km2，是太湖流域较大的湖泊，具有蓄水、养殖、供水、航运、旅游等多种社会和生态功能。最近20多年来，滆湖流域工农业发展极为迅速，成为中国最具经济活力的地区之一。然而伴随着经济的繁荣，进入滆湖的营养负荷逐年增加，滆湖的渔业捕捞强度不断攀升，滆湖生态环境遭到极大破坏。
鲢鳙是我国传统水产养殖的主要种类，也是世界上引种最多的种类之一。自从非经典生物操纵治理蓝藻水华的理论提出以后，对鲢鳙在湖泊中的生态地位的研究逐渐增多，多家研究单位的多个研究人员通过围隔、室内受控水体等对鲢鳙滤食藻类、对氮磷循环的影响等进行了研究。本论文利用ecopath模型，比较分析了1986-1989和2010年两个时期滆湖生态系统的特征。并在滆湖敞水区建立了2050亩的围网养殖区域，研究鲢鳙对湖泊水质、浮游植物、底栖动物和沉积物中氮、磷的影响。
1滆湖生态系统的ecopath模型研究
利用1986～1989年的生态调查和渔业统计数据，构建了一个具有21个功能组的ecopath模型，利用软件自带的基本分析（Basicestimates）和网络分析（Networkanalysis）功能，对1986-1989年滆湖生态系统的结构和功能进行了量化和系统分析。对模型分析结果表明，1986-1989年滆湖生态系统包含7个营养级，主要的营养物质流动发生在前4个营养级。功能组的营养级从1～3.584。系统总流量为12131.76t·km~(-2)·year-1，其中第I营养级的流量占总流量的74.91%。系统总净初级生产力为3722.117t·km~(-2)·year-1，其中水草生产量占总生产量的33.31%。平均捕捞营养级为2.78，大部分渔业目标群体（青鱼、草鱼、中华绒螯蟹、鲢、鳙、鲤、团头鲂等）的EE（EcotrophicEfficience，EE）都较高，表明这些鱼类种群增长受到渔业和捕食的联合作用限制。P/R（Production/Respiratory，P/R）值为2.76，P/B（Production/Biomass，P/B）值为1.76，Finn's循环指数为14.76%，依据Odum对成熟生态系统特征的描述，作者认为滆湖在1980年代后期是相对成熟的生态系统。
利用2010年滆湖的生态调查和渔业统计数据，构建了滆湖在2010年生态系统的ecopath模型。模型包含17个功能组，对模型进行分析的结果表明，2010年滆湖生态系统包含7个营养级，各功能组的营养级范围为1～3.691，其营养物质流动主要发生在前4个营养级；系统总生产力为1974.82t·km~(-2)·year-1，总流量为8562.544t·km~(-2)·year-1，联结指数、系统杂食系数、Finn’s循环指数以及Finn’s平均路径长度分别为0.219、0.189、7.99%和2.841；P/R、P/B分别为2.189和3.509；平均捕捞营养级为2.56。通过对1986-1989年和2010年滆湖ecopath生态模型的总流量、P/R、P/B、Finn′s循环指数、联接指数、食物网优势食物链等特征的比较可知，和1986-1989年相比，2010年的滆湖生态系统处于从成熟期退化的过程中，渔业资源衰退，系统中K-对策者比例减小而r-对策者比例增加，即渔业也是湖泊富营养化的受害者。根据混合营养效应分析，鲢鳙现有生物量不能有效控制滆湖藻类的生物量。
2围网放养鲢鳙对水质和浮游植物的影响
在滆湖南部支河口以西2km处建立了面积为2050亩的围网，分为3个实验组，放养鲢的密度分别为194、206和242尾/亩，放养鳙的密度分别为133、148和186尾/亩。在围网内外共设置了17个监测点，从2009年12月至2010年12月每月一次对浮游生物、水质、底栖动物和沉积物理化特征进行监测分析。结果表明，放养鲢鳙可以有效降低围网内的叶绿素含量，而对于总磷、总氮和氨氮来说，围网内数值稍低于围网外，但是差异不显著。
对于浮游植物，围网内密度显著低于围网外，而生物量则是围网内稍低于围网外，差异不显著。围网内不同鲢鳙密度对叶绿素a含量没有显著影响。围网外优势种主要有铜绿微囊藻、微小色球藻、梅尼小环藻、小空星藻和四尾栅藻等，围网内主要有铜绿微囊藻、微小色球藻、针状蓝纤维藻、尖尾蓝隐藻、二形栅藻等；即优势种以蓝藻门、绿藻门和硅藻门种类为主，围网内外优势种没有明显区别。
3围网放养鲢鳙对底栖动物的影响
鲢鳙对底栖动物的影响主要是通过改变营养循环、影响底栖动物生存环境等起作用。通过近1年的监测，在围网区域共发现底栖动物16种，分属于环节动物门、节肢动物门和软体动物门。其中环节动物门最多，有11种，占67%；节肢动物门次之，有4种，占28%；软体动物门仅有1种。霍甫水丝蚓、铜锈环棱螺、苏氏尾鳃蚓为围网养殖区域的优势种。围网外底栖动物密度显著高于围网内，生物量则是围网内稍高于围网外。优势种基本相同，围网外皮氏管水蚓（Aulodrilus pigueti）重要值更高。
4围网放养鲢鳙对沉积物中氮磷的影响
围网内沉积物中总氮低于围网外，但差异不显著；围网内沉积物中总氮呈上升趋势，而围网外沉积物中总氮则呈下降趋势。围网内沉积物中总磷显著高于围网外，且内外都表现出下降的趋势，但是围网内沉积物中总磷下降更快。鲢鳙对沉积物中氮磷表现出一种复杂的影响，表面上看来，放养鲢鳙会导致沉积物中总磷含量减少（相对于围网外，围网内沉积物中的总磷含量下降得更快），而总氮含量增加。结合围网内水层氮磷比逐渐增高，那么一个可能的原因是从化学计量学的角度来解释，即鲢鳙作为氮、磷的汇，将更多的磷固定在鱼体内，并且促进了沉积物中磷的释放。
虽然本次研究中围网规模和全湖相比仍然较小，加上湖流的影响，鲢鳙难以对整个湖泊产生深刻的影响，鲢鳙对富营养化湖泊的影响还需要更大规模的实验数据支持，但本论文对于理解鲢鳙在湖泊营养循环中的作用机制是一个有益的探索。
Lake Gehu is the secondary big freshwater lake in South Jiangsu province, China.With the area of164km2, Lake Gehu is relative lager in watershed of Lake Taihu andplay multiple roles of social and ecological such as irrigation, aquaculture, water supply,shipping and tourism. Development of agriculture and industry in Lake Gehu watershedwas so quick and Lake Gehu has becoming one of the most vigorous area in China inthe past two decades. Otherwise more and more nutrients loading entered Lake Gehuand fisheries density continue increasing, accompanied economic booming, made LakeGehu suffered serious ecological damage.
To find out reasons of deteriation of Lake Gehu ecology and the recovery ways,resolve problems of retreat of fisheries resource and degradation of environment, manyinstitutes investigated the environment, geograph, ecology, fisheries and biology andaccumulated plenty of foundamental data. During “eleven five” plan, many sub-projectsof state water pollutant control and handle programm were related to Lake Gehu andplenty of work has made. But most of these researches was carried out from one aspectof ecosystem and lake of systematic decision and behavior under the frame ofecosystem architecture. Computer model of food web based on mass balance couldanalysis ecosystem holistic and integrate basic research data and documents. Domesticscholars have conducted a preliminary study using computer models on the ecosystemof Taihu Lake and East China Sea. In this thesis we constructed a EwE model on behalfof the Gehu ecosystem history and current situation by synthesizing theory and methodsfrom biology, ecology, ecological stoichiometry and computer science, analysized andquantified the food web composition, distribution of trophic levels, ecosystem maturityand stability, influence of fisheries policy. Results would future guiding fisheries policydevelopment and ecological restoration of Gehu. This contribution first constructedmodel of Gehu and made a comparison with model in1986-1989and found that Gehuwas under degradation compare to1986-1989.
Silver and bighead carp are major kindes in china traditional aquaculture and oneof the most introduced fishes in the world. Since the theory of non-classicalbiomanipulation of control cyanobacterial blooms was put forward, researches aboutniches of silver carp and bighead carp in the lakes is gradually increased, a number of researchers in different institutes studied filtration on algae and influence on nitrogenand phosphorus cycling of silver and bighead carp through enclosure, indoor controlledexperinments. The researchers draw different and even opposite conclusions for theyused different experimental methods and in different environments. About140hectarepen culture with structure improved set up in the open area of Gehu, stocking differentdensity of silver carp and bighead carp, to verify influence that the silver carp andbighead carp in natural water bodies on the lake ecology, provide guidance on controlcyanobacterial blooms and eutrophication recovery.
1Ecopath model study of Lake Gehu ecosystem
An ecopath model with21functional groups was constructed based on the data ofecological investigation and fisheries statistics of1986-1989. Structure andcharacteristics of Gehu ecosystem was analysized by network analysis moduleembedded in ecopath. The analysis results demonstrated that ecosystem in1986-1989included seven trophic levels and major trophic flow occurred in the first four levels.Range of groups from within1-3.584. Total system throughput was12131.76t·km~(-2)·year-1, among this flow originated from first trophic level accounted for74.91%.Total net system primary production was3722.117t·km~(-2)·year-1, among thismacrophyte production accounted for33.31%. Mean catched trophic level was2.78.Ecotrophic efficience of most target fishes were high, indicated that most of these fisheswere constrained by combination of fisheries and predatory. Value of P/R(Production/Respiratory, P/R) was2.76, value of P/B (Production/Biomass, P/B) was1.76, Finn's cycling index was14.76%. According criteria from Odum of matureecosystem, we considered that Gehu ecosystem in1986-1989was mature.
Another ecopath model of Lake Gehu on2010was constructed using data ofinvestigation and fisheries statistics. The model included seventeen groups. Analysisresults indicate that ecosystem of2010Gehu Lake comprised of seven trophic level.Range of trophic level of these groups was1-3.691and major flow occurred in the firstfour trophic level. Total system production was1974.82t·km~(-2)·year-1, total systemthroughput was8562.544t·km~(-2)·year-1, connectance index, system omnivore index,Finn’s cycling index and Finn’s mean path length were0.219,0.189,7.99and2.841respectively. Values of P/R and P/B were2.189and3.509respectively. Mean catchedtrophic level was2.56. Through comparison of total system throughput, P/R, P/B,Finn’s cycling index, connectance index and dominant food chain between1986-1989and2010, we could conclude that Gehu was degrading from mature status, resouces offisheries declined, K-strategists decreasing and r-strategists increasing, present biomassof silver and bighead carp could not control biomass of algae.
2Experinment study of influence of silver carp and bighead carp onphotoplankton
Pen culture was set up in the west of Zhihekou, south of Gehu, with the area ofabout140hectares. Three treatments with different density of silver and bighead carpwas carried out. Density of silver carp were2910,3090and3630/hm2respectively anddensity of bighead carp were1995,2220and2775/hm2respectively. Seventeensampling sites were configured outside and inside the pen culture. Investgatedplankton, water quality, zoobenthos and nitrogen and phosphorus in sediment montyly.The analysis results showed that stocking of silver and bighead carp could decreasingchlorophyll a significantly, but could not decrease total nitrogen, togal phosphorus andammonia nitrogen significantly. Density of phytoplankton ouside of the pen culturewas significant greater than that of inside the pen culture but there is no significantdifference of biomass betweeen inside and outside of the pen culture.
3Influence of silver carp and bighead carp on zoobenthos
Silver and bighead carp influenced zoobenthos mainly through altering nutrientscycling and modifying habitat of zoobenthos. We found16zoobenthos in the area ofpen culture within studied period and they belonged to Annelida, Arthropoda andMollusca. Among these zoobenthos11belonged to annelida,67%;4belonged toArthropoda,28%; and1belonged to Mollusca. Limnodrilus hoffmeisteri, Aulodriluspigueti and Bellamya aeruginosa are dominant species in the pen culture area.
4Influence of silver carp and bighead carp on nitrogen and phosphorus insediment
Total nitrogen inside the pen culture was lower than that of outside of thepenculture but there was no significant difference. Total phosphorus inside of the penculture was significant greater than that of outside the pen culture. The conclusion wasthat stock of silver carp and bighead carp could decrease phosphorus accumulation inthe sediment and had no vital influence on nitroten.
This contribution was a positive explore to understand the mechanism that silvercarp and bighead carp played in the lake nutrients cycling. Limitatios of this study wasthat the area of pen culture was small compare to the whole lake, with the interere of lake currence the silver carp and bighead carp could influenced the whole lake profund.Research on this field still need data come from bigger scale.
鲢鳙是我国传统水产养殖的主要种类，也是世界上引种最多的种类之一。自从非经典生物操纵治理蓝藻水华的理论提出以后，对鲢鳙在湖泊中的生态地位的研究逐渐增多，多家研究单位的多个研究人员通过围隔、室内受控水体等对鲢鳙滤食藻类、对氮磷循环的影响等进行了研究。本论文利用ecopath模型，比较分析了1986-1989和2010年两个时期滆湖生态系统的特征。并在滆湖敞水区建立了2050亩的围网养殖区域，研究鲢鳙对湖泊水质、浮游植物、底栖动物和沉积物中氮、磷的影响。
1滆湖生态系统的ecopath模型研究
利用1986～1989年的生态调查和渔业统计数据，构建了一个具有21个功能组的ecopath模型，利用软件自带的基本分析（Basicestimates）和网络分析（Networkanalysis）功能，对1986-1989年滆湖生态系统的结构和功能进行了量化和系统分析。对模型分析结果表明，1986-1989年滆湖生态系统包含7个营养级，主要的营养物质流动发生在前4个营养级。功能组的营养级从1～3.584。系统总流量为12131.76t·km~(-2)·year-1，其中第I营养级的流量占总流量的74.91%。系统总净初级生产力为3722.117t·km~(-2)·year-1，其中水草生产量占总生产量的33.31%。平均捕捞营养级为2.78，大部分渔业目标群体（青鱼、草鱼、中华绒螯蟹、鲢、鳙、鲤、团头鲂等）的EE（EcotrophicEfficience，EE）都较高，表明这些鱼类种群增长受到渔业和捕食的联合作用限制。P/R（Production/Respiratory，P/R）值为2.76，P/B（Production/Biomass，P/B）值为1.76，Finn's循环指数为14.76%，依据Odum对成熟生态系统特征的描述，作者认为滆湖在1980年代后期是相对成熟的生态系统。
利用2010年滆湖的生态调查和渔业统计数据，构建了滆湖在2010年生态系统的ecopath模型。模型包含17个功能组，对模型进行分析的结果表明，2010年滆湖生态系统包含7个营养级，各功能组的营养级范围为1～3.691，其营养物质流动主要发生在前4个营养级；系统总生产力为1974.82t·km~(-2)·year-1，总流量为8562.544t·km~(-2)·year-1，联结指数、系统杂食系数、Finn’s循环指数以及Finn’s平均路径长度分别为0.219、0.189、7.99%和2.841；P/R、P/B分别为2.189和3.509；平均捕捞营养级为2.56。通过对1986-1989年和2010年滆湖ecopath生态模型的总流量、P/R、P/B、Finn′s循环指数、联接指数、食物网优势食物链等特征的比较可知，和1986-1989年相比，2010年的滆湖生态系统处于从成熟期退化的过程中，渔业资源衰退，系统中K-对策者比例减小而r-对策者比例增加，即渔业也是湖泊富营养化的受害者。根据混合营养效应分析，鲢鳙现有生物量不能有效控制滆湖藻类的生物量。
2围网放养鲢鳙对水质和浮游植物的影响
在滆湖南部支河口以西2km处建立了面积为2050亩的围网，分为3个实验组，放养鲢的密度分别为194、206和242尾/亩，放养鳙的密度分别为133、148和186尾/亩。在围网内外共设置了17个监测点，从2009年12月至2010年12月每月一次对浮游生物、水质、底栖动物和沉积物理化特征进行监测分析。结果表明，放养鲢鳙可以有效降低围网内的叶绿素含量，而对于总磷、总氮和氨氮来说，围网内数值稍低于围网外，但是差异不显著。
对于浮游植物，围网内密度显著低于围网外，而生物量则是围网内稍低于围网外，差异不显著。围网内不同鲢鳙密度对叶绿素a含量没有显著影响。围网外优势种主要有铜绿微囊藻、微小色球藻、梅尼小环藻、小空星藻和四尾栅藻等，围网内主要有铜绿微囊藻、微小色球藻、针状蓝纤维藻、尖尾蓝隐藻、二形栅藻等；即优势种以蓝藻门、绿藻门和硅藻门种类为主，围网内外优势种没有明显区别。
3围网放养鲢鳙对底栖动物的影响
鲢鳙对底栖动物的影响主要是通过改变营养循环、影响底栖动物生存环境等起作用。通过近1年的监测，在围网区域共发现底栖动物16种，分属于环节动物门、节肢动物门和软体动物门。其中环节动物门最多，有11种，占67%；节肢动物门次之，有4种，占28%；软体动物门仅有1种。霍甫水丝蚓、铜锈环棱螺、苏氏尾鳃蚓为围网养殖区域的优势种。围网外底栖动物密度显著高于围网内，生物量则是围网内稍高于围网外。优势种基本相同，围网外皮氏管水蚓（Aulodrilus pigueti）重要值更高。
4围网放养鲢鳙对沉积物中氮磷的影响
围网内沉积物中总氮低于围网外，但差异不显著；围网内沉积物中总氮呈上升趋势，而围网外沉积物中总氮则呈下降趋势。围网内沉积物中总磷显著高于围网外，且内外都表现出下降的趋势，但是围网内沉积物中总磷下降更快。鲢鳙对沉积物中氮磷表现出一种复杂的影响，表面上看来，放养鲢鳙会导致沉积物中总磷含量减少（相对于围网外，围网内沉积物中的总磷含量下降得更快），而总氮含量增加。结合围网内水层氮磷比逐渐增高，那么一个可能的原因是从化学计量学的角度来解释，即鲢鳙作为氮、磷的汇，将更多的磷固定在鱼体内，并且促进了沉积物中磷的释放。
虽然本次研究中围网规模和全湖相比仍然较小，加上湖流的影响，鲢鳙难以对整个湖泊产生深刻的影响，鲢鳙对富营养化湖泊的影响还需要更大规模的实验数据支持，但本论文对于理解鲢鳙在湖泊营养循环中的作用机制是一个有益的探索。
Lake Gehu is the secondary big freshwater lake in South Jiangsu province, China.With the area of164km2, Lake Gehu is relative lager in watershed of Lake Taihu andplay multiple roles of social and ecological such as irrigation, aquaculture, water supply,shipping and tourism. Development of agriculture and industry in Lake Gehu watershedwas so quick and Lake Gehu has becoming one of the most vigorous area in China inthe past two decades. Otherwise more and more nutrients loading entered Lake Gehuand fisheries density continue increasing, accompanied economic booming, made LakeGehu suffered serious ecological damage.
To find out reasons of deteriation of Lake Gehu ecology and the recovery ways,resolve problems of retreat of fisheries resource and degradation of environment, manyinstitutes investigated the environment, geograph, ecology, fisheries and biology andaccumulated plenty of foundamental data. During “eleven five” plan, many sub-projectsof state water pollutant control and handle programm were related to Lake Gehu andplenty of work has made. But most of these researches was carried out from one aspectof ecosystem and lake of systematic decision and behavior under the frame ofecosystem architecture. Computer model of food web based on mass balance couldanalysis ecosystem holistic and integrate basic research data and documents. Domesticscholars have conducted a preliminary study using computer models on the ecosystemof Taihu Lake and East China Sea. In this thesis we constructed a EwE model on behalfof the Gehu ecosystem history and current situation by synthesizing theory and methodsfrom biology, ecology, ecological stoichiometry and computer science, analysized andquantified the food web composition, distribution of trophic levels, ecosystem maturityand stability, influence of fisheries policy. Results would future guiding fisheries policydevelopment and ecological restoration of Gehu. This contribution first constructedmodel of Gehu and made a comparison with model in1986-1989and found that Gehuwas under degradation compare to1986-1989.
Silver and bighead carp are major kindes in china traditional aquaculture and oneof the most introduced fishes in the world. Since the theory of non-classicalbiomanipulation of control cyanobacterial blooms was put forward, researches aboutniches of silver carp and bighead carp in the lakes is gradually increased, a number of researchers in different institutes studied filtration on algae and influence on nitrogenand phosphorus cycling of silver and bighead carp through enclosure, indoor controlledexperinments. The researchers draw different and even opposite conclusions for theyused different experimental methods and in different environments. About140hectarepen culture with structure improved set up in the open area of Gehu, stocking differentdensity of silver carp and bighead carp, to verify influence that the silver carp andbighead carp in natural water bodies on the lake ecology, provide guidance on controlcyanobacterial blooms and eutrophication recovery.
1Ecopath model study of Lake Gehu ecosystem
An ecopath model with21functional groups was constructed based on the data ofecological investigation and fisheries statistics of1986-1989. Structure andcharacteristics of Gehu ecosystem was analysized by network analysis moduleembedded in ecopath. The analysis results demonstrated that ecosystem in1986-1989included seven trophic levels and major trophic flow occurred in the first four levels.Range of groups from within1-3.584. Total system throughput was12131.76t·km~(-2)·year-1, among this flow originated from first trophic level accounted for74.91%.Total net system primary production was3722.117t·km~(-2)·year-1, among thismacrophyte production accounted for33.31%. Mean catched trophic level was2.78.Ecotrophic efficience of most target fishes were high, indicated that most of these fisheswere constrained by combination of fisheries and predatory. Value of P/R(Production/Respiratory, P/R) was2.76, value of P/B (Production/Biomass, P/B) was1.76, Finn's cycling index was14.76%. According criteria from Odum of matureecosystem, we considered that Gehu ecosystem in1986-1989was mature.
Another ecopath model of Lake Gehu on2010was constructed using data ofinvestigation and fisheries statistics. The model included seventeen groups. Analysisresults indicate that ecosystem of2010Gehu Lake comprised of seven trophic level.Range of trophic level of these groups was1-3.691and major flow occurred in the firstfour trophic level. Total system production was1974.82t·km~(-2)·year-1, total systemthroughput was8562.544t·km~(-2)·year-1, connectance index, system omnivore index,Finn’s cycling index and Finn’s mean path length were0.219,0.189,7.99and2.841respectively. Values of P/R and P/B were2.189and3.509respectively. Mean catchedtrophic level was2.56. Through comparison of total system throughput, P/R, P/B,Finn’s cycling index, connectance index and dominant food chain between1986-1989and2010, we could conclude that Gehu was degrading from mature status, resouces offisheries declined, K-strategists decreasing and r-strategists increasing, present biomassof silver and bighead carp could not control biomass of algae.
2Experinment study of influence of silver carp and bighead carp onphotoplankton
Pen culture was set up in the west of Zhihekou, south of Gehu, with the area ofabout140hectares. Three treatments with different density of silver and bighead carpwas carried out. Density of silver carp were2910,3090and3630/hm2respectively anddensity of bighead carp were1995,2220and2775/hm2respectively. Seventeensampling sites were configured outside and inside the pen culture. Investgatedplankton, water quality, zoobenthos and nitrogen and phosphorus in sediment montyly.The analysis results showed that stocking of silver and bighead carp could decreasingchlorophyll a significantly, but could not decrease total nitrogen, togal phosphorus andammonia nitrogen significantly. Density of phytoplankton ouside of the pen culturewas significant greater than that of inside the pen culture but there is no significantdifference of biomass betweeen inside and outside of the pen culture.
3Influence of silver carp and bighead carp on zoobenthos
Silver and bighead carp influenced zoobenthos mainly through altering nutrientscycling and modifying habitat of zoobenthos. We found16zoobenthos in the area ofpen culture within studied period and they belonged to Annelida, Arthropoda andMollusca. Among these zoobenthos11belonged to annelida,67%;4belonged toArthropoda,28%; and1belonged to Mollusca. Limnodrilus hoffmeisteri, Aulodriluspigueti and Bellamya aeruginosa are dominant species in the pen culture area.
4Influence of silver carp and bighead carp on nitrogen and phosphorus insediment
Total nitrogen inside the pen culture was lower than that of outside of thepenculture but there was no significant difference. Total phosphorus inside of the penculture was significant greater than that of outside the pen culture. The conclusion wasthat stock of silver carp and bighead carp could decrease phosphorus accumulation inthe sediment and had no vital influence on nitroten.
This contribution was a positive explore to understand the mechanism that silvercarp and bighead carp played in the lake nutrients cycling. Limitatios of this study wasthat the area of pen culture was small compare to the whole lake, with the interere of lake currence the silver carp and bighead carp could influenced the whole lake profund.Research on this field still need data come from bigger scale.
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