Paradox of nutrient removal in coupled socioeconomic and ecological dynamics for lake water pollution

We study coupled socioeconomical and ecological dynamics for lake water pollution. Players choose between cooperative (but costly) option and economical option, and their decision is affected by the fraction of cooperators in the community and by the importance of water pollution problem. When an opportunity for choice arrives, players take the option with the higher utility (best response dynamics). This social dynamics is coupled with the dynamics of lake water pollution. First, oscillation of large amplitude is generated if social change occurs faster than ecosystem responses. Second, the model can show “paradox of nutrient removal”. If phosphorus is removed more effectively either from the inflow or from the lake water, the pollution level may increase (rather than decrease) due to the decline in people's willingness to cooperate. Third, we compare the effectiveness of alternative methods in improving water quality: to reduce the cooperation cost by subsidy, to enhance people's concern to water pollution problem, and to promote the conformity among people.     

Humans copy rapidly increasing choices in a multiarmed bandit problem

Conformist social learning, the tendency to acquire the most common trait in a group, allows individuals to rapidly acquire established beneficial traits from a multitude of options. However, conformist strategies hinder acquisition of novel advantageous behavior patterns, because such innovations are by definition uncommon. This raises the possibility that proxy cues of the success of novel traits may be utilized to identify and acquire advantageous innovations and disregard failing options. We show that humans use changes in trait frequency over time as such a cue in an economic game. Participants played a three-alternative forced choice game (i.e., a multi-armed bandit), using social information to attempt to locate a high reward that could change location. Participants viewed temporal changes in how many players chose each option in two successive rounds. Participants supplemented conformist strategies with a “copy-increasing-traits” strategy. That is, regardless of the traits absolute population frequencies, participants' choices were guided by changes in trait frequencies. Thus, humans can detect advantageous innovations by monitoring how many individuals adopt these over time, adopting traits increasing in frequency, and abandoning traits decreasing in frequency. Copying rapidly increasing traits allows identification and acquisition of advantageous innovations, and is thus potentially key in facilitating their early diffusion and cultural evolution.     

Effects of Conformism on the Cultural Evolution of Social Behaviour

Models of cultural evolution study how the distribution of cultural traits changes over time. The dynamics of cultural evolution strongly depends on the way these traits are transmitted between individuals by social learning. Two prominent forms of social learning are payoff-based learning (imitating others that have higher payoffs) and conformist learning (imitating locally common behaviours). How payoff-based and conformist learning affect the cultural evolution of cooperation is currently a matter of lively debate, but few studies systematically analyse the interplay of these forms of social learning. Here we perform such a study by investigating how the interaction of payoff-based and conformist learning affects the outcome of cultural evolution in three social contexts. First, we develop a simple argument that provides insights into how the outcome of cultural evolution will change when more and more conformist learning is added to payoff-based learning. In a social dilemma (e.g. a Prisoner’s Dilemma), conformism can turn cooperation into a stable equilibrium; in an evasion game (e.g. a Hawk-Dove game or a Snowdrift game) conformism tends to destabilize the polymorphic equilibrium; and in a coordination game (e.g. a Stag Hunt game), conformism changes the basin of attraction of the two equilibria. Second, we analyse a stochastic event-based model, revealing that conformism increases the speed of cultural evolution towards pure equilibria. Individual-based simulations as well as the analysis of the diffusion approximation of the stochastic model by and large confirm our findings. Third, we investigate the effect of an increasing degree of conformism on cultural group selection in a group-structured population. We conclude that, in contrast to statements in the literature, conformism hinders rather than promotes the evolution of cooperation.     

茅尾海营养状况及其来源研究

根据2008年茅尾海海域水质调查结果,采用营养指数法、营养状态质量指数法、有机污染指数法对海湾营养状况进行了评价,并结合2007年入海污染源调查结果,探讨了茅尾海的营养盐来源。结果表明:茅尾海受无机氮污染较重,海域处于呈富营养状态,有机污染程度属2级,表明开始受到有机污染。营养盐高值区集中在北部海域和东部海域。钦江、茅岭江等入海河流携带入海的营养盐是海域营养的主要来源,占入海污染物总量的79%以上,其次来自钦州湾外海的混合排污口。2001~2007年茅尾海DIN、DIP年均浓度分别与入海河流氮、磷营养盐入海负荷呈显著正相关,两者的相关系数分别为0.873和0.824。     

Analyzing the mutual feedbacks between lake pollution and human behaviour in a mathematical social-ecological model

Does the adoption of environment-oriented actions by individuals necessarily improve the state of an ecosystem in the most effective way? We address this question with the example of eutrophication in shallow lakes. When exposed to fertilizers, such lakes can undergo a critical transition called eutrophication, resulting in a loss of biodiversity and ecosystem services. We couple a generic model of eutrophication with a best-response model of human behaviour, where agents can choose to pollute the lake at a high level (defection) or at a low level (cooperation). It is known that feedbacks between the interacting lake pollution and human behaviour can give rise to complex dynamics with multiple stable states and oscillations. Here, we analyze the impact of all model parameters on the shape of the nullclines. S-shaped nullclines are a condition for complex dynamics to occur. Moreover, we find that agents decreasing their pollution discharge into the lake is not necessarily the most effective way to reduce the pollution level in the lake. This is due to coexisting counterintuitive stable equilibria where the lake is in a clear state despite a high level of pollution discharge. We analyze the complex dynamics of the system and describe in detail Hopf, saddle-node, homoclinic and Bogdanov-Takens bifurcations. The complex dynamics with potential multistability and counterintuitive equilibria suggests that generic management recommendations holding for every level of pollution and of cooperation are impossible. Apart from the direct perturbation of an ecological variable, we identify three ways a management strategy can influence the social-ecological system: it can change the location, the resilience and the existence of stable equilibria.     

Do descriptive social norms drive peer punishment? Conditional punishment strategies and their impact on cooperation

Peer punishment is widely considered a key mechanism supporting cooperation in human groups. Although much research shows that human behavior is shaped by the prevailing social norms, little is known about how punishment decisions are impacted by the social context. We present a set of large-scale incentivized experiments in which participants (999 American participants recruited via Amazon Mechanical Turk) could punish their partner conditional on either the level of cooperation or the level of punishment displayed by others who previously interacted in the same setting. While many participants punish independently of levels of cooperation or punishment, a substantial portion punishes free riding more severely when cooperation is more common (‘norm enforcement’), or when free riding is more severely punished by others (‘conformist punishment’). With a dynamic model we demonstrate that conditional punishment strategies can substantially promote cooperation. In particular, conformist punishment helps cooperation to gain a foothold in a population, and norm enforcement helps to maintain cooperation at high levels. Our results provide solid empirical evidence of conditional punishment strategies and illustrate their possible implications for the dynamics of human cooperation.     

Why people punish defectors. Weak conformist transmission can stabilize costly enforcement of norms in cooperative dilemmas

In this paper, we present a cultural evolutionary model in which norms for cooperation and punishment are acquired via two cognitive mechanisms: (1) payoff-biased transmission-a tendency to copy the most successful individual; and (2) conformist transmission-a tendency to copy the most frequent behavior in the population. We first show that if a finite number of punishment stages is permitted (e.g. two stages of punishment occur if some individuals punish people who fail to punish non-cooperators), then an arbitrarily small amount of conformist transmission will stabilize cooperative behavior by stabilizing punishment at some n -th stage. We then explain how, once cooperation is stabilized in one group, it may spread through a multi-group population via cultural group selection. Finally, once cooperation is prevalent, we show how prosocial genes favoring cooperation and punishment may invade in the wake of cultural group selection.     

厦门市海岸带水污染负荷估算及预测

综合采用灰色模型、曲线回归等预测方法,建立了基于厦门市海岸带特征的主要污染源水污染负荷估算及预测模型,并采用厦门市历年统计数据对模型加以验证.对厦门市近岸海域近10年的废水和主要污染物质排放量估算的结果表明:万元产值工业废水排放量呈逐年下降的趋势,而各污染物的排放总量却逐年缓慢增长;在点源污水排放总量预测中,约76%的氮、磷来自于生活污水;在非点源污染负荷中,农业非点源中的氮、磷负荷占较大比例,城市非点源污染负荷比例最小.2005年厦门海岸带各污染源产生的氮污染负荷大小比较结果为:生活污染源>农业非点源>工业污染源>旅游业污染源>城市非点源,磷污染负荷则为:农业非点源>生活污染源>工业污染源>旅游业污染源>城市非点源.     

Restoration and resilience to recovery of the Lake Loosdrecht ecosystem in relation to its phosphorus flow

A reduction in external phosphorus loading since 1984 to Loosdrecht lakes system by the dephosphorization of the inlet water, yielded only minor effects in Lake Loosdrecht. This reduction measure turned out to have decreased the loading only by a factor of two. A conceptual model was constructed based on laboratory measurements to describe phosphorus flow in the lake ecosystem for the summer of 1987. The role of zooplankton and fish was more important in phosphorus recycling than diffusion at the sediment-water interface. The input and output of phosphorus of the lake were at equilibrium and therefore, further reduction in external loading was needed for recovery. The results of the conceptual model agreed well with the output of the mathematical model PCLOOS. Additional measures such as dredging, flushing, chemomanipulation, or biomanipulation would be ineffective at the present level of external loading. Only a significant further reduction in external input will restore Lake Loosdrecht's water quality over a long period of time.     

Social learning in cooperative dilemmas

Helping is a cornerstone of social organization and commonplace in human societies. A major challenge for the evolutionary sciences is to explain how cooperation is maintained in large populations with high levels of migration, conditions under which cooperators can be exploited by selfish individuals. Cultural group selection models posit that such large-scale cooperation evolves via selection acting on populations among which behavioural variation is maintained by the cultural transmission of cooperative norms. These models assume that individuals acquire cooperative strategies via social learning. This assumption remains empirically untested. Here, I test this by investigating whether individuals employ conformist or payoff-biased learning in public goods games conducted in 14 villages of a forager–horticulturist society, the Pahari Korwa of India. Individuals did not show a clear tendency to conform or to be payoff-biased and are highly variable in their use of social learning. This variation is partly explained by both individual and village characteristics. The tendency to conform decreases and to be payoff-biased increases as the value of the modal contribution increases. These findings suggest that the use of social learning in cooperative dilemmas is contingent on individuals'' circumstances and environments, and question the existence of stably transmitted cultural norms of cooperation.     

Assessing the role of internal phosphorus recycling on eutrophication in four lakes in China and Malaysia

Internal phosphorus recycling (IPR) is an important nutrient source driving algal growth and eutrophication in lakes. The complexity of eutrophication behaviours caused by high IPR complicates lake management and undermines restoration efforts. Hence, knowledge about the possible types of bifurcation behaviours caused by high IPR is essential for effective and sustainable lake eutrophication management. For this purpose, numerical bifurcation analysis is performed on an algae‑phosphorus model to investigate how IPR drives complex and rich eutrophication behaviours in two tropical and two subtropical lakes. The two tropical lakes are Tasik Harapan and Sunway Lagoon in Malaysia, while the two subtropical lakes are Lake Fuxian and Lake Taihu in China. For each specified level of IPR, co-dimension one bifurcation analysis is performed by means of XPPAUT. Co-dimension two bifurcation analysis is then carried out by means of MatCont. At low IPR, Lake Fuxian exhibits reversible behaviour, accompanied by higher external phosphorus loading (EPL) thresholds. Lake Fuxian is also more conducive to stable equilibrium and its lake dynamics are easily predictable. At moderate IPR, Sunway Lagoon is likely to exhibit stable equilibrium, accompanied by possible shifting between two stable steady states (hysteresis behaviour) and oscillations. With higher IPR, Lake Taihu and Tasik Harapan are prone to irreversibility, accompanied by lower EPL thresholds. Because of increased complexity in lake dynamics in Lake Taihu and Tasik Harapan, small changes in EPL or in algal mortality rates could trigger various transitions in lake dynamics. Overall, high IPR can trigger unexpected sharp increases in algal concentration and can reduce the resilience of an oligotrophic lake. For shallow lakes, high IPR would cause unexpected sharp increases in algal concentrations, undermine resilience of lakes, complicate lake management, and delay lake recovery process.     

Why cooperation is not running away

A growing number of experimental and theoretical studies show the importance of partner choice as a mechanism to promote the evolution of cooperation, especially in humans. In this paper, we focus on the question of the precise quantitative level of cooperation that should evolve under this mechanism. When individuals compete to be chosen by others, their level of investment in cooperation evolves towards higher values, a process called competitive altruism, or runaway cooperation. Using a classic adaptive dynamics model, we first show that when the cost of changing partner is low, this runaway process can lead to a profitless escalation of cooperation. In the extreme, when partner choice is entirely frictionless, cooperation even increases up to a level where its cost entirely cancels out its benefit. That is, at evolutionary equilibrium, individuals gain the same payoff than if they had not cooperated at all. Second, importing models from matching theory in economics we, however, show that when individuals can plastically modulate their choosiness in function of their own cooperation level, partner choice stops being a runaway competition to outbid others and becomes a competition to form the most optimal partnerships. In this case, when the cost of changing partner tends towards zero, partner choice leads to the evolution of the socially optimum level of cooperation. This last result could explain the observation that human cooperation seems to be often constrained by considerations of social efficiency.     

The evolution of social learning rules: Payoff-biased and frequency-dependent biased transmission

Humans and other animals do not use social learning indiscriminately, rather, natural selection has favoured the evolution of social learning rules that make selective use of social learning to acquire relevant information in a changing environment. We present a gene-culture coevolutionary analysis of a small selection of such rules (unbiased social learning, payoff-biased social learning and frequency-dependent biased social learning, including conformism and anti-conformism) in a population of asocial learners where the environment is subject to a constant probability of change to a novel state. We define conditions under which each rule evolves to a genetically polymorphic equilibrium. We find that payoff-biased social learning may evolve under high levels of environmental variation if the fitness benefit associated with the acquired behaviour is either high or low but not of intermediate value. In contrast, both conformist and anti-conformist biases can become fixed when environment variation is low, whereupon the mean fitness in the population is higher than for a population of asocial learners. Our examination of the population dynamics reveals stable limit cycles under conformist and anti-conformist biases and some highly complex dynamics including chaos. Anti-conformists can out-compete conformists when conditions favour a low equilibrium frequency of the learned behaviour. We conclude that evolution, punctuated by the repeated successful invasion of different social learning rules, should continuously favour a reduction in the equilibrium frequency of asocial learning, and propose that, among competing social learning rules, the dominant rule will be the one that can persist with the lowest frequency of asocial learning.     

Whitemere, a lake that defies some conventions about nutrients

1. Whitemere, a kettle-hole lake in north-west England (z_m,14 m, area 22.5 ha) has extremely high maximal concentrations (around 1 mg L⁻¹) of total phosphorus (TP) and soluble reactive phosphorus (SRP), and comparatively low maximal concentrations of dissolved inorganic nitrogen (<0.5 mg N L⁻¹). Bioassays indicate that its phytoplankton is nitrogen limited, and it has surface blue-green algal blooms in summer. Palaeoecological investigations have shown that high populations of cyanophytes occurred more than 6000 years ago, even before human settlement of the catchment.
2. The reasons for this combination of features are not immediately clear. This paper attempts to distinguish between two hypotheses: that the high phosphorus concentrations arise from input of phosphate-rich groundwater; and that the high phosphorus concentrations arise from concentration and recycling mechanisms within the lake.
3. The lake is entirely fed by ground water, direct rainfall and surface sheet flow. The concentrations of phosphorus in the ground water are much lower than in the lake water. The lake is stratified and has high phosphorus concentrations in the anaerobic hypolimnion in summer as a result of substantial release from the sediment. Vertical turbulence in the relatively weakly stratified water column may transfer substantial amounts to the epilimnion.There is also substantial phosphorus release from the sediments under the aerobic epilimnion.
4. There is little outflow of water and phosphorus. Most of the phosphorus is returned to the sediment under winter isothermal conditions, to be released again the following summer. The lake has probably maintained these mechanisms for millennia. Recent eutrophication is likely to have been driven by nitrogen inputs from greater intensification of agriculture in the catchment.     

The spatial distribution of nutrients in the South Basin of Windermere

SUMMARY. The vertical and horizontal distribution of electrical conductivity, soluble reactive phosphorus (SRP), total phosphorus, ammonia-nitrogen and nitrate-nitrogen in the South Basin of Windermere is described and related, where possible, to spatial variations in phytoplankton and zooplankton. For most variables, the maximum range of horizontal variation on a single day was greater than the maximum range of seasonal variation measured at a single station over a full year. Horizontal variations in SRP and ammonia were particularly high, with coefficients of variation often exceeding 100%. The errors associated with sampling at a single station were most pronounced when local accumulations of zooplankton or phytoplankton occurred in the lake. Horizontal variations in conductivity were primarily influenced by the discharge of treated sewage effluent into the central region and the mass transport of water from the more oligotrophic North Basin. Occasionally, more widespread variations in conductivity appeared to be related to spatial variations in photosynthetic activity. The major factor influencing the horizontal distribution of SRP was the discharge of treated effluent into the central region. Despite the intensive recycling of phosphorus, SRP concentrations were nearly always 10–30% higher near the sewage works than elsewhere in the basin. On a few occasions, significant horizontal differences in SRP concentration were also detected within downwind accumulations of crustacean zooplankton. Horizontal variations in total phosphorus were produced by spatial differences in SRP concentration or by local accumulations of phytoplankton or zooplankton. Horizontal differences in ammonia concentration appeared to be produced, both by the effluent discharge and by the turbulent transfer of nutrients from sediments in shallow water. Observations of effluent movement and dispersion demonstrate that wind-induced water movements tend to recirculate nutrient-rich water in the central region and limit mixing along the axis of the basin. The statistical implications of nutrient heterogeneity are discussed in relation to sampling strategy and the possible effects of persistent nutrient concentration gradients on phytoplankton patch formation are assessed.     

Microcommunities and microgradients: Linking nutrient regeneration,microbial mutualism,and high sustained aquatic primary production

Nutrient regeneration is essential to sustained primary production in the aquatic environment because of coupled physical and metabolic gradients. The commonly evaluated ecosystem perspective of nutrient regeneration, as is illustrated among planktonic paradigms of lake ecosystems, functions only at macrotemporal and spatial scales. Most inland waters are small and shallow. Consequently, most organic matter of these waters is derived from photosynthesis of emergent, floating-leaved, and submersed higher plants and microflora associated with living substrata and detritus, including sediments, as well as terrestrial sources. The dominant primary productivity of inland aquatic ecosystems is not planktonic, but rather is associated with surfaces. The high sustained rates of primary production among sessile communities are possible because of the intensive internal recycling of nutrients, including carbon. Steep gradients exist within these attached microbial communities that (a) require rapid, intensive recycling of carbon, phosphorus, nitrogen, and other nutrients between producers, particulate and dissolved detritus, and bacteria and protists: (b) augment internal community recycling and losses with small external inputs of carbon and nutrients from the overlying water or from the supporting substrata; and (c) encourage maximal conservation of nutrients. Examples of microenvironmental recycling of carbon, phosphorus, and oxygen among epiphytic, epipelic, and epilithic communities are explained. Recalcitrant dissolved organic compounds from decomposition can serve both as carbon and energy substrates as well as be selectively inhibitory to microbial metabolism and nutrient recycling. Rapid recycling of nutrient and organic carbon within micro-environments operates at all levels, planktonic as well as attached, and is mandatory for high sustained productivity.     

应用原生动物群落评价枯水期白洋淀的水质现状

应用PFU微型生物群落监测方法,对长期处于低水位状态下的白洋淀水质进行了综合评价。结果表明,白洋淀水域受府河污水的影响,淀内的内源性污染,对原生动物群落造成了极大的影响。与20世纪90年代的资料相比,淀内中心区4个采样点原生动物的群集种类数(1d)急剧下降,而丰度上升了1．28倍,表明内源性污染对白洋淀水质的影响加剧。     

武汉东湖沉积物的环境地球化学

武汉东湖沉积物的地球化学过程受到人类活动的较大影响。本文报道该湖沉积物的沉积性状和ＣＮ、Ｐ、Ｓｉ、Ｃａ、Ｍｇ、Ｆｅ、Ａｌ、Ｍｎ、Ｃｕ、Ｐｂ、Ｚｎ、Ｃｄ诸元素在沉积层序中的分布,研究了这些元素的地球化学。通过对东湖污染状况和类型的分析,指出营养物质的积累是导致湖泊生态环境恶化的重要原因,提出可用磷含量作为划分东湖沉积物污染程度的指标的观点。     

应用多种生物类群评价白洋淀水环境质量变化

白洋淀是华北平原最大的浅水湖泊。近几十年来,白洋淀上游耗水增加,天然入淀水量急剧减少,使白洋淀水位明显降低;同时,随着淀泊周边农村生活污水的排入,白洋淀发生了严重的富营养化。2017年雄安新区设立,白洋淀环境治理工作的重要性空前提高。近几年通过上游水库和跨流域生态补水,白洋淀水位明显抬升,同时淀中和淀边农村实施了生活污水收集。为了解白洋淀当前水环境状况,对前期补水和治污的效果进行评估,2019年8月对白洋淀水体理化指标及水生生物进行调查,采用水质、浮游植物、浮游动物、底栖动物指标评价白洋淀水环境质量,并与2010年的调查数据进行对比,辅以沉水植物指标评价水体生态状况。结果表明: 白洋淀2019年总磷达到优于Ⅲ类水平,总氮优于Ⅱ类水平,水环境状况比2010年有较大改善。其中,总磷浓度下降了88.6%,总氮浓度下降了83.9%,叶绿素a浓度下降了47.8%,透明度增大了43.4%;浮游动物、底栖动物等生物多样性明显提高,浮游植物密度显著降低,优势种从普遍的重富营养型转变为富营养型;沉水植物清洁型种类分布范围缩小。这说明沉水植物物种评价法不适于白洋淀水环境质量评价,浮游动物多样性评价方法也不适用于当前白洋淀水环境质量评价。早期,由于污染源集中在西部,白洋淀西部水质差于东部;现在,由于自西、北、南多方向补水,淀区的水动力条件发生显著改变,加之西部实施治污工程,淀区水质呈现空间均一化的特点。     

磷模型在千岛湖水体污染预测中的应用研究

运用国际上通用的多个经验统计磷模型预测千岛湖目前的磷浓度水平,其中第6个模型最适合用于千岛湖的预测预报．利用该模型预测了磷负荷量分别减少1/4、1/2、3/4情况下,千岛湖水体中总磷浓度的响应．同时针对千岛湖营养水平控制的短、长期目标,利用模型计算出了所允许的总磷负荷量及入湖磷浓度．