热带典型珊瑚岛礁海洋牧场花刺参底播增殖容量及其生态效应预测

为改善热带珊瑚岛礁型海洋牧场的珊瑚礁生境,实现生物资源的养护和渔业资源的产出功能,在对海参等高值经济种开展底播增殖前,科学评估其生态容量是防止引发海洋牧场生态风险的重要保证。运用生态系统模型法评估了三亚蜈支洲岛热带珊瑚岛礁海洋牧场花刺参(Stichopus monotuberculatus)的底播增殖容量。根据2020~2021年蜈支洲岛海洋牧场近岛区渔业资源调查与环境因子数据,运用Ecopath with Ecosim 6.6软件构建了该海域的生态系统营养通道模型。研究表明:生态系统各功能组营养级范围介于1~3.52,系统的食物网结构以牧食食物链为主,总能流中有43%的能量来源于碎屑功能组,其在系统总能流中有重要地位。系统的总平均能量传递效率为9.353%,略低于林德曼能量传递效率(10%)。总初级生产量/总呼吸量为3.726,总初级生产量/总生物量为28.834,系统连接指数为0.256,杂食性指数为0.120,系统Finn''s循环指数和平均路径长度分别为2.485%和2.379,表明近岛区生态系统食物网结构较为简单,且系统稳定性和成熟度偏低,易受外界干扰。根据模型评估的花刺参增殖生态容量为110.21 t/km²,是现存量的206 倍,有较大增殖空间,并且达到生态容量后碎屑组的能量再循环利用效率将显著增加,营养级结构能得到进一步优化,系统稳定性及成熟度将有所提高。基于研究结果,可适当采捕与花刺参生态位相近的生物,同时增殖放流其他处于不同营养层次的经济种,从而减少种间竞争,有效利用系统冗余能量,进而扩大花刺参的生态容量,实现海洋牧场的健康可持续发展。     

Ecosystem modelling in the southern Benguela: comparisons of Atlantis,Ecopath with Ecosim,and OSMOSE under fishing scenarios

Ecosystem-based management of marine fisheries requires the use of simulation modelling to investigate the system-level impact of candidate fisheries management strategies. However, testing of fundamental assumptions such as system structure or process formulations is rarely done. In this study, we compare the output of three different ecosystem models (Atlantis, Ecopath with Ecosim, and OSMOSE) applied to the same ecosystem (the southern Benguela), to explore which ecosystem effects of fishing are most sensitive to model uncertainty. We subjected the models to two contrasting fishing pressure scenarios, applying high fishing pressure to either small pelagic fish or to adult hake. We compared the resulting model behaviour at a system level, and also at the level of model groups. We analysed the outputs in terms of various commonly used ecosystem indicators, and found some similarities in the overall behaviour of the models, despite major differences in model formulation and assumptions. Direction of change in system-level indicators was consistent for all models under the hake pressure scenario, although discrepancies emerged under the small-pelagic-fish scenario. Studying biomass response of individual model groups was key to understanding more integrated system-level metrics. All three models are based on existing knowledge of the system, and the convergence of model results increases confidence in the robustness of the model outputs. Points of divergence in the model results suggest important areas of future study. The use of feeding guilds to provide indicators for fish species at an aggregated level was explored, and proved to be an interesting alternative to aggregation by trophic level.     

Quantitative model of trophic interactions in Beibu Gulf ecosystem in the northern South China Sea

1IntroductionThe Beibu Gulf is a natural semiclosed conti-nental sea of the South China Sea,which is situatedat17°00′~21°45′N,105°40′~110°10′E,and sur-rounded by China and Vietnam(see Fig.1).It hasa subtropic monsoon climate with an average winter     

Food web and fish stock changes in central Chile: comparing the roles of jumbo squid (Dosidicus gigas) predation,the environment,and fisheries

We analyzed recent food web and fish stock changes in the central Chile marine ecosystem, comparing the roles of jumbo squid (Dosidicus gigas) as predator, the environment, and fishing. To accomplish this we used food web modeling and the Ecopath with Ecosim software (EwE). The principal fish stocks have experienced wide decadal fluctuations in the past 30 years, including stock collapses of horse mackerel (Trachurus murphyi) and hake (Merluccius gayi), and there was a large influx of jumbo squid during the mid-2000s. We used two EwE models representing the food web off central Chile to test the hypothesis that predation by jumbo squid has been significant in explaining the dynamics of the main fishing resources and other species in the study area. Results indicate that predation by jumbo squid on fish stocks is lower than that of other predators (e.g. hake) and the fishery. Long-term fluctuations (1978–2004) in the biomass of the main fish stocks (as well as other components of the food web) seem to be related to fishing and to variation in primary production, rather than to predation by jumbo squid alone. Jumbo squid seems to play a role as predator rather than prey in the system, but its impacts are low when compared with the impacts of other predators and fishing. Therefore, we conclude that jumbo squid predation on its prey was not the primary force behind the collapse of important fish stocks off central Chile. Future efforts should be directed to better understanding factors that trigger sudden increases in jumbo squid abundance off central Chile, as well as modeling its trophic impacts.     

A trophic model of the Pearl River Delta coastal ecosystem

A 26-compartment steady-state trophic model (1997–1999) was constructed using the Ecopath with Ecosim software to study the general status and development trends of the Pearl River Delta coastal ecosystem. The results show that the values of effective trophic level ranged from 1.00 to 4.21. It was found that a high trophic niche overlap existed in the typical estuarine ecosystem. Mixed trophic impacts show that detritus and the groups at the low trophic levels had positive influences on most groups. The ecosystem was found to be in an immature state during 1997–1999 based on the system statistics.     

Changes in trophic flows and ecosystem properties of the Beibu Gulf ecosystem before and after the collapse of fish stocks

Mass-balance models (Ecopath) of the ecosystem before and after collapse (1959-1961 and 1997-1999) of fish stocks were developed with Ecopath software to compare the differences in ecosystem structure, functioning and ecosystem properties of the Beibu Gulf. The model includes 20 functional groups consisting of commercial important fish groups and other ecologically important groups in the ecosystem such as zooplankton, phytoplankton, and detritus. Results indicated that biomass and catches of the system have changed drastically between the 1960s and 1990s, especially for the high trophic levels (TL). The biomass of level V in the early 1960s was 32 times higher than that of the late 1990s, however, the biomass of level I and II in the 1990s was higher than the 1960s. Despite the higher catches in the 1990s, fishing was ecologically less expensive during the 1990s than 1960s due to small fish catches were large. Mean transfer efficiency decreased from for 10.2% in the 1960s to 9.1% in the 1990s periods. According to the summary statistics, the parameters of net system production (NPS) and total primary production to total respiration ratio were increased from 1.013 in the 1960s to 2.184 in the 1990s, however, the connectance index (CI), system omnivore index, Finn’s cycling index and mean path length decreased from the 1960s to the 1990s. The overhead (O) was higher in the 1990s model while the ascendancy (A) decreased nearly 10% in the 1960s. The ‘Keystoneness’ result indicate that zooplankton was identified as keystone species in 1960s, however, the elasmobranches was keystone species in the late 1990s. The average trophic level of the fishery decreased from 3.32 in the 1960s to 2.98 in the 1990s, and exhibits classic symptoms of “fishing down the food web”. All the indices of the system attributes suggests that the Beibu Gulf ecosystem in 1960s was found to be more mature than in the 1990s due to the collapse of demersal ecosystem, and the ecosystem changed from being dominated by long-lived, high trophic level groundfish dominated system toward a system with small-size and low-value species over fifty years.     

A spatio-temporal ecosystem model to simulate fishing management plans: A case of study in the Gulf of Gabes (Tunisia)

The Gulf of Gabes located in southern Tunisia is one of the most productive ecosystems in the Mediterranean Sea. Despite its ecological importance, it is subject to high fishing pressure affecting the different components of the ecosystem. Given the multispecies, multigear nature of the fishery, there is a need to manage trade-offs between environmental and economic objectives. In this study, an Ecospace model was developed based on the previously constructed Ecopath model of the Gulf of Gabes and calibrated for the period 1995–2008 to investigate the response of the ecosystem to a set of alternative spatial management scenarios. These scenarios were derived from the current fishery regulation owing the important interest expressed by local fishery managers to assess new management measures. The results showed for each management scenario how bottom trawling and coastal fishing impact the different trophic groups and the complexity of interaction between these two fishing activities. Furthermore, spatially explicit simulations were performed to identify regions where the management measures are effective. Results suggested that for some trophic groups, these regions are well-defined which would be interesting to propose more accurate spatial measures. Finally, several indicators were calculated to evaluate the proposed management plans and provide managers with a straightforward set of decision rules to describe the potential trade-offs and fulfill both fisheries and conservation management objectives in the context of an ecosystem approach. The decision rules were based on observed trends to reduce uncertainty relative to the model complexity and provide consistent advice to decision-makers.     

基于Ecopath模型的海州湾及邻近海域生态系统结构研究

基于2018年海州湾及邻近海域的渔业资源底拖网调查数据,运用Ecopath with Ecosim 6.5 (EwE)软件构建由26个功能群组成的海州湾及邻近海域生态系统Ecopath模型,对现阶段该生态系统的营养结构、营养相互关系和系统总特征等进行分析,旨在为实施基于生态系统的渔业管理提供理论依据。结果表明:海州湾及邻近海域生态系统各功能群的营养级范围为1.00～4.19,其中鱼类营养级范围较广,为3.22～4.19;浮游动物和其他软体动物受初级生产者和捕食者的双重作用,处于重要的营养位置;生态系统总体特征分析显示,该生态系统的总初级生产量与总呼吸量的比值为7.096,总初级生产量与总生物量的比值为56.866,系统的连接指数和系统杂食指数分别为0.429和0.204,说明该生态系统目前处于不成熟、不稳定的状态,容易受外界扰动的影响。本文通过对海州湾及邻近海域生态系统模型进行研究,解析了该海域营养结构和系统发育状况,将为海州湾渔业资源的可持续利用和科学管理提供理论依据。     

Construction and analysis of a coral reef trophic network for Qilianyu Islands,Xisha Islands

Qilianyu Islands coral reefs (QICR), located in the northeastern part of the South China Sea, has been affected by human activities and natural disturbance. To characterize the trophic structure, ecosystem properties and keystone species of this region, a food-web model for the QICR is developed using methods involving a mass-balance approach with Ecopath with Ecosim software. Trophic levels range from 1.00 for detritus and primary producers to 3.80 for chondrichthyes. The mean trophic transfer efficiency for the entire ecosystem is 13.15%, with 55% of total energy flow originating from primary producers. A mixed trophic impact analysis indicates that coral strongly impacts most components of this ecosystem. A comparison of our QICR model with that for other coral reef ecosystems suggests that the QICR ecosystem is immature and/or is degraded.     

应用物质平衡模型模拟一个半封闭海湾的营养结构和能流特征

The marine ecosystem of the Jiaozhou Bay has degraded significantly in fisheries productivity and its ecological roles as spawning and nursery ground for many species of commercial importance has been declining in recent years. A mass-balanced trophic model was developed using Ecopath with Ecosim to evaluate the trophic structure of the Jiaozhou Bay for improving ecosystem management. The model were parameterized based on the fisheries survey data in the Jiaozhou Bay in 2011, including 23 species groups and one detritus group according to their ecological roles. The trophic levels of these ecological groups ranged from 1(primary producers and detritus) to4.3(large demersal fishes). The estimated total system throughput was 12 917.10 t/(km~2·a), with 74.59% and25.41% contribution of the total energy flows from phytoplankton and detritus, respectively. Network analyses showed that the overall transfer efficiency of the ecosystem was 14.4%, and the mean transfer efficiency was 14.5%for grazing food chain and 13.9% for detritus food chain. The system omnivory index(SOI), Finn's cycled index(FCI) and connectance index(CI) were relatively low in this area while the total primary production/total respiration(TPP/TR) was high, indicating an immature and unstable status of the Jiaozhou Bay ecosystem. Mixed trophic impact analysis revealed that the cultured shellfish had substantial negative impacts on most functional groups. This study contributed to ecosystem-level evaluation and management planning of the Jiaozhou Bay ecosystem.     

Changes in the trophic structure of the southern Benguela before and after the onset of industrial fishing

Despite a human presence in the Benguela region for at least one million years, exploitation of marine resources by European seafarers only began in earnest in the 1400s. Ecopath with Ecosim was used to construct and compare mass-balanced foodweb models of the southern Benguela ecosystem, representing the following eras of human influence: aboriginal (10 000 BP–1651), pre-industrial (1652–1909), industrial (1910–1974) and post-industrial (1975–present). Biomass at higher trophic levels (TLs) decreased over the periods examined, whereas that of sardine and anchovy increased in the early 2000s, reflected by the decline in weighted TL of the community (excluding plankton). Fishing became an important predatory impact, taking over consumption of small pelagics and horse mackerel from declined natural predators such as hake. Harvesting of apex predators such as seals and seabirds during the pre-industrial era meant that the mean TL of the catch declined markedly between the pre-industrial (1900) and industrial (1960) models. Biomass removals by fishing have increased substantially over time. Total biomass, consumption, respiration, production and throughput decreased from the pristine model to 1960 and then increased again in the 2000s, probably influenced by the abnormally high small pelagic biomass in the early 2000s. Three additional alternate scenarios were examined for each of the retrospective models, in particular to explore the effects of removing large fish and forage fish from the system. Although biomasses and consumption of various groups in these scenarios differed from base models, indicators such as TL of the community and piscivore groups, and the diversity indices, were not altered much, suggesting that outputs from such retrospective models in the form of derived, relative indicators, may be more robust than comparisons of absolute flows, although the latter provide supplementary inferences. Although South African fisheries have certainly impacted ecosystem structure since their commencement, these effects are in addition to natural (specifically environmental) forcing that has always been influencing the system. Fishing stress at the ecosystem level and the collapse of small pelagic stocks may lead to a shift toward a bottom-up trophic control mechanism becoming the dominant driver of ecosystem dynamics, increasing the impact of environmental events including climate change. It is thus possible that pristine systems were not as severely affected by environmental anomalies as are modern systems.     

基于增殖放流的定栖性种类剩余产量模型及其模拟分析

为应对渔业资源的日益衰退,增殖放流成为了目前补充资源、维持资源可持续利用的主要手段之一。增殖放流实施后,渔业资源的可持续特征是学者们普遍关心,却又无法使用传统剩余产量模型有效解决的问题。本研究基于传统的Schaefer剩余产量模型,提出了一个适用于增殖放流情况下的剩余产量模型(增殖剩余产量模型),模拟分析了不同增殖放流和捕捞策略对模型的影响。该模型的形式与Schaefer剩余产量模型相似,但加入了描述增殖群体增长特征的参数—有效增殖率,以此来表示增殖放流的群体对产量产生的影响。结果显示,合理的增殖放流可以起到增加最大可持续产量的效果,使用增殖剩余产量模型能够得到合理的最大可持续产量等关键指标的估算结果。与无增殖放流情况相比,在增殖放流影响下,海域原存资源(海域原本存在的群体)达到最大可持续产量时所需的生物量较小,而可承受的捕捞努力量则有所增加。增殖剩余产量模型所反映的原存群体和增殖群体之间会产生抑制作用。在该作用影响下,不同增殖放流和捕捞策略会对模型的评估结果产生影响。与传统模型相比,该模型将增殖放流纳入最大可持续产量的评估过程,提高了增殖放流影响下最大可持续产量评估的准确性,可用于诸如海洋牧场等边界较清晰的海域内增殖定栖性种类最大可持续产量的估算。     

Trophic network model of the Northern Adriatic Sea: Analysis of an exploited and eutrophic ecosystem

A quantitative model of the trophic network of Northern Adriatic Sea marine ecosystem during the 1990s has been constructed, with the goal of analysing its trophic structure, identifying the key trophic groups and assessing the anthropogenic impacts on the ecosystem using the Ecopath modelling protocol. The Northern Adriatic Sea is an eutrophic, shallow basin, and one of the most heavily fished areas in the Mediterranean Sea. The network aggregation into discrete trophic levels sensu Lindeman shows that low trophic levels dominate biomass and energy flows, with 40% of the total system throughput flowing out from trophic level 2. Instead, upper trophic levels appear bottom-up controlled, highly depleted and not exerting any control on the trophic network, as shown by mixed trophic impact-based analyses. Microbial loop is comparable to grazing with respect to the magnitude of flows involved, as 66% of the trophic network flows originate from detritus, which is mainly consumed by bacteria. Key trophic groups are plankton groups, macro-crustaceans and detritus, and other r-selected organisms like squids and small pelagics, which have a great influence on the ecosystem. In particular, zooplankton acts as a bottleneck for energy flows, limiting the energy from the low trophic levels effectively reaching the upper food web. The high pelagic production caused by eutrophication sustains high fishery landings and impressive discard quantities, as well as the benthic compartment. Overall, the ecosystem appears quite productive and in a stressed and developmental status. Model results and comparisons with few existing historical data suggest that the low maturity and stressed state of the Northern Adriatic Sea are not only due to natural characteristics, but mainly to anthropogenic pressures.     

An evaluation of underlying mechanisms for “fishing down marine food webs”

自1998年“捕捞降低海洋食物网”概念首次提出以来,渔获物平均营养级（Mean trophic level of fisheries landings,MTL）广泛用于评估捕捞活动对生态系统完整性的影响,并指导管理机构的政策制定。近年来研究表明,掌握MTL的潜在变化机制对于以MTL作为渔业可持续性指标至关重要。根据联合国粮农组织FAO提供的渔获统计数据,结合Fishbase提供的相关鱼种营养级,本文探讨了全球三大洋14个FAO渔区MTL的变化趋势,并进一步分析不同MTL变化趋势下高营养级鱼种和低营养级鱼种渔获量的变动情况。研究表明, MTL呈上升和回升状态均可能伴随着低营养级鱼种渔获量的下降。此外,通过观察营养级高于3.25鱼种的MTL以区分“捕捞降低海洋食物网”和“捕捞沿着海洋食物网”现象需考虑生态系统的群落结构和开发历程。利用渔获物平均营养级评价渔业可持续性必须综合考虑高营养级和低营养级鱼种,以及群落结构和开发历程的掩盖效应。     

三亚蜈支洲岛海洋牧场近岛区底表大型底栖动物群落结构及评价

三亚蜈支洲岛海洋牧场拥有丰富的岛礁生物资源。为掌握海洋牧场近岛区的底表大型底栖动物群落组成和分布特征及其影响因子、评估海洋牧场底栖生态系统健康状况, 本研究于2020—2021年进行了底表大型无脊椎动物群落季节变动的调查。结果表明: 近岛珊瑚礁区共鉴定出棘皮动物、软体动物、节肢动物3大门类90种, 其中秋季种类数最多为55种, 夏季种类数最少为16种; 底表大型底栖动物的年平均丰度为0.87±0.26ind.·m^-2, 年平均生物量为76.99±34.32g·m^-2。群落聚类分析(cluster)与多维排序尺度分析(multidimensional Scaling, MDS)表明, 该区域群落结构季节性差异不显著, 各站位间群落结构受沉积物性质以及人类活动的频繁程度的影响, 形成北部与南部区域2个聚类组。全年的物种丰富度指数d为2.7±1.16, 多样性指数H′为3.14±0.88, 均匀度指数J为0.76±0.11。基于多样性指数H'及多变量海洋生物指数(Multivariate-AZTI′s marine biotic index, M-AMBI)评价指标, 海洋牧场近岛区环境除夏季为轻度污染外, 其他季节均为无污染状态。采用动物丰度与生物量比较曲线(abundance-biomass curves, ABC曲线)法评价底栖动物群落稳定性状况得出, 除冬季以外, 其他季节底表大型底栖动物群落受到一定程度干扰, 尤其是夏季群落结构稳定性较低。建议应持续关注底表大型底栖动物群落变动, 调整、优化涉海休闲旅游活动, 以保证蜈支洲岛海洋牧场生态系统的长期健康与稳定。     

Evaluating impacts of pulse fishing on the effectiveness of seasonal closure

Seasonal fishing closures are often used in fisheries management to conserve overfished stocks.As one of the unintended consequences,fishermen often contend for maximizing catches immediately after reopening fisheries.The resultant large catch landings in a short time period(i.e.,pulse fishing)may undermine the benefit of closure.We implemented an end-to-end model OSMOSE-JZB(Object-oriented Simulator of Marine ec OSystem Exploitation OSMOSE)modelling ecosystem in the Jiaozhou Bay located in China to evaluate the impact of pulse fishing on the effectiveness of seasonal closure at levels of fish community,population,and individual.Our study demonstrated that the three-month closure was successful in conserving fish stocks.There were small variations on ecological indicators(i.e.,total biomass of the community,mean trophic level of the community,mean trophic level of the catch,and Shannon-Wiener biodiversity index)when pulse fishing occurred.Pulse fishing seemed not to result in a great shift in community structure.Compared to other species,the biomass of two large predatory fishes were more susceptible to pulse fishing.Pulse fishing could change the pressure of predators to fish stocks via food webs,especially for young individuals.Our simulations indicate that we can improve the effectiveness of seasonal closure by managing pulse fishing.Although the results derived in this study may be specific to the target ecosystem,the general approach is applicable to other ecosystems when evaluating fishing impacts.     

Interactions between a natural food web,shellfish farming and exotic species: The case of the Bay of Mont Saint Michel (France)

To ensure sustainable uses of the coastal zone, an integrated ecosystemic approach and ecosystem models are required to frame ecological processes and evaluate environmental impacts. Here, a mass-balance trophic (Ecopath) model of the Mont Saint Michel Bay (MSMB) was developed, to analyze the bay's functioning as an ecosystem. This bay, intensively exploited by fishing and for shellfish farming, is also suffering from the proliferation of the gastropod Crepidula fornicata, an exotic species.     

现代海洋牧场建设之我见

现代海洋牧场建设是实现我国海洋渔业与近海生态系统和谐发展的重要途径之一,作为一种海洋经济新业态,海洋牧场既能养护渔业资源,又能修复生态环境。本文论述了现代海洋牧场建设的必要性,分析了我国海洋牧场建设中存在的建设技术缺乏标准化、承载力评估缺乏模型化、建设理念缺乏生态化、管理缺乏信息化等问题,探讨了现代海洋牧场相关理念及分类模式。针对现代海洋牧场建设存在的突出问题,提出了构建陆海统筹现代海洋牧场亟待解决的技术先导和"四化"突破(工程化、机械化、自动化和信息化)等对策和建议。     

基于Ecopath模型的七连屿礁栖性生物的生态承载力分析

生态承载力评估是开展生物资源增殖放流, 维持珊瑚礁生态系统健康的基础和前提。本文基于2019年渔业资源和生态环境的综合调查数据, 构建了七连屿珊瑚礁海域生态系统的生态通道(Ecopath)模型, 分析和探讨了相关功能组增殖放流的生态承载力。结果显示, 七连屿珊瑚礁海域生态系统各功能群营养级范围为1.00~3.81; 生态系统的总能量转化效率为13.45%; 生态系统以牧食食物链占据主导地位, 直接来源于初级生产者的能流占比为57%。系统总初级生产量/总呼吸量为2.54; 总初级生产量/总生物量为19.07; 系统连接指数和系统杂食性指数分别为0.36和0.22, 表明当前七连屿珊瑚礁海域生态系统的成熟度和稳定性偏低, 系统对于外界的干扰抵抗能力较弱。在未改变七连屿珊瑚礁生态系统结构和功能的前提下, 各功能组中珊瑚、双壳类和植食性鱼类的生态承载力分别为25.09~53.77t•km^-2、2.55~39.95t•km^-2和4.89~17.94t•km^-2, 因此仍具有较大的增殖空间。珊瑚礁鱼类群落的最大生态承载力同珊瑚礁无脊椎动物群落的增殖密切相关, 在未来的珊瑚礁渔业管理中应从生态系统整体结构的角度综合考虑增殖放流的方法设计。     

Comparing pristine and depleted ecosystems: The Sørfjord, Norway versus the Gulf of St. Lawrence, Canada. Effects of intense fisheries on marine ecosystems

The Sørfjord, Norway, and the Gulf of St. Lawrence, Canada, are two sub-arctic ecosystems with similar trophic structure. However, in the Gulf of St. Lawrence, severe exploitation of groundfish stocks has lead to important shifts in the trophic structure. In the Sørfjord, the situation is different: fishing pressure is much lighter. Our hypothesis is that overexploitation leads to changes in the trophic structure and severely alters the resilience of ecosystems. Based on the same modelling approach (Ecopath with Ecosim) the food web structure was compared, using different ecosystem indicators. Patterns of food web structure and trophodynamics were contrasted. Cod was the keystone species in both ecosystems, and forage fish were also important. Even after similar environmental changes in both ecosystems, and after a reduction of fishing pressure in the Gulf of St. Lawrence, there is no recovery of cod stocks in this ecosystem. In the Sørfjord, after different perturbations (but not from the fishery), the ecosystem seems to return to its equilibrium.