Open systems living in a closed biosphere: a new paradox for the Gaia debate

While energetically open, the biosphere is appreciably closed from the standpoint of matter exchange. Matter cycling and recycling is hence a necessary and emergent property of the global-scale system known as Gaia. But how can an aggregate of open-system life forms have evolved and persisted for billions of years within a planetary system that is largely closed to matter influx and outflow? The puzzling nature of a closed yet persistent biosphere draws our attention to the course of evolution of fundamental metabolic strategies and matter-capture techniques. It suggests a facet of the Gaia hypothesis, framed in terms of persistence. The oceans, atmosphere, soils and biota constitute a complex system which maintains and adjusts matter cycling and recycling within the constraints of planetary closure such that open-system forms of life can persist. This weaker version of the Gaia hypothesis may be useful because it readily lends itself to at least one form of test. What is the solution to the closed biosphere puzzle, and does it indicate that Gaia merits status as a discrete entity? We suggest several disciplines within the field of biology that might provide tools and perspectives toward reaching a solution. These disciplines include artificial closed ecosystems, prokaryote evolution, the nexus of thermodynamics and evolutionary biology, and hierarchy theory in ecosystem modeling and evolution theory.     

The fundamental processes in ecology: a thought experiment on extraterrestrial biospheres

Ecological science is often organised as a hierarchical series of entities: genes, individuals, populations, species, communities, ecosystems and biosphere. Here, I consider an alternative process‐based approach to ecology, and analyse the nature of the fundamental processes in ecology. These fundamental processes are discussed in the context of the following question:‘for any planet with carbon‐based life, which persists over geological time scales, what are the minimum set of ecological processes that must be present?‘I suggest that the following processes would be present on any such planet: energy flow, multiple guilds, ecological tradeo ffs leading to within‐guild biodiversity, ecological hypercycles, merging of organismal and ecological physiology, carbon sequestration and possibly photosynthesis. Nutrient cycling is described as an emergent property of these fundamental processes. I discuss reasons why a biosphere based on a single species with no nutrient cycling is very unlikely to exist. I also describe the concept of ‘Gaian effect’. This suggests that some processes will always tend to extend the lifespan of a biosphere in which they develop (positive Gaian effect) while others could either increase or decrease (negative Gaian effect) such a lifespan. These ideas are discussed in the context of astrobiology, ecosystem services, conservation biology and Gaia theory.     

The SER Standards,cultural ecosystems,and the nature‐culture nexus—a reply to Evans and Davis

Evans and Davis claim the SER Standards use a “pure naturalness” model for restoration baselines and exclude most cultural ecosystems from the ecological restoration paradigm. The SER Standards do neither. The SER Standards consider both “natural” ecosystems (that are unequivocally not cultural) and “similar” cultural ecosystems as suitable reference models. Furthermore, Evans and Davis propose assessing whether a cultural ecosystem exhibits “good, bad, or neutral impacts from humans on ecosystems” as the basis for reference models. We argue that such an approach would overlook the indispensability of native ecosystem benchmarks to measure human impacts and provide a springboard for social‐ecological restoration.     

Coral chimerism as an evolutionary rescue mechanism to mitigate global climate change impacts

Climate change and anthropogenic pressures inflict a wide range of profound damages on coral reef ecosystems, reshaping coral reef communities due to their physiological and ecological intolerance to the newly developing environmental conditions. Here, I present coral chimerism as an evolutionary rescue tool for accelerating adaptive responses to global climate change impacts. The “evolutionary rescue” power is contingent on the premise that coral chimerism counters the erosion of genetic and phenotypic diversity. Further benefits are gained when flexible chimeric entities alter their somatic constituents following changes in environmental conditions, synergistically presenting the best‐fitting combination of their genetic components to endure in a capricious environment, exhibiting always their environmentally matched physiological characteristics. Chimerism should be considered as an integral part of the ecological engineering toolbox being developed for active reef restoration.     

Natural selection through survival alone,and the possibility of Gaia

Here I advance two related evolutionary propositions. (1) Natural selection is most often considered to require competition between reproducing “individuals”, sometimes quite broadly conceived, as in cases of clonal, species or multispecies-community selection. But differential survival of non-competing and non-reproducing individuals will also result in increasing frequencies of survival-promoting “adaptations” among survivors, and thus is also a kind of natural selection. (2) Darwinists have challenged the view that the Earth’s biosphere is an evolved global homeostatic system. Since there is only one biosphere, reproductive competition cannot have been involved in selection for such survival-promoting adaptations, they claim. But natural selection through survival could reconcile Gaia with evolutionary theory.     

Indispensable role of quantum theory in the brain dynamics

Recently, Tegmark pointed out that the superposition of ion states involved in the superposition of firing and resting states of a neuron quickly decohere. It undoubtedly indicates that neural networks cannot work as quantum computers, or computers taking advantage of coherent states. Does it also mean that the brain can be modeled as a neural network obeying classical physics? Here we show that it does not mean that the brain can be modeled as a neural network obeying classical physics. A brand new perspective in research of neural networks from quantum theoretical aspect is presented.     

Warming weakens facilitative interactions between decomposers and detritivores,and modifies freshwater ecosystem functioning

Warming is among the major drivers of changes in biotic interactions and, in turn, ecosystem functioning. The decomposition process occurs in a chain of facilitative interactions between detritivores and microorganisms. It remains unclear, however, what effect warming may have on the interrelations between detritivores and microorganisms, and the consequences for the functioning of natural freshwater ecosystems. To address these gaps, we performed a field experiment using tank bromeliads and their associated aquatic fauna. We manipulated the presence of bacteria and detritivorous macroinvertebrates (control, “bacteria,” and “bacteria + macroinvertebrates”) under ambient and warming scenarios, and analyzed the effects on the microorganisms and ecosystem functioning (detritus mass loss, colored dissolved organic matter, and nitrogen flux). We applied antibiotic solution to eliminate or reduce bacteria from control bromeliads. After 60 days incubation, bacterial density was higher in the presence than in the absence of macroinvertebrates. In the absence of macroinvertebrates, temperature did not influence bacterial density. However, in the presence of macroinvertebrates, bacterial density decreased by 54% with warming. The magnitude of the effects of organisms on ecosystem functioning was higher in the combined presence of bacteria and macroinvertebrates. However, warming reduced the overall positive effects of detritivores on bacterial density, which in turn, cascaded down to ecosystem functioning by decreasing decomposition and nitrogen flux. These results show the existence of facilitative mechanisms between bacteria and detritivores in the decomposition process, which might collapse due to warming. Detritivores seem to contribute to nutrient cycling as they facilitate bacterial populations, probably by increasing nutrient input (feces) in the ecosystem. However, increased temperature mitigated these beneficial effects. Our results add to a growing research body that shows that warming can affect the structure of aquatic communities, and highlight the importance of considering the interactive effects between facilitation and climatic drivers on the functioning of freshwater ecosystems.     

长江口滨海湿地潮间带生态系统的多稳态特征

多稳态现象普遍存在于多种生态系统中,它与生态系统的健康和可持续发展密切相关,已成为生态学研究的热点与难点,但是目前有关滨海湿地生态系统多稳态的形成机制还缺乏深入研究.本文以崇明东滩鸟类自然保护区的潮间带生态系统为研究对象,通过以下内容,开展滨海湿地多稳态研究: 1)通过验证多稳态的判定依据“双峰”和“阈值”特征,证实长江口潮间带生态系统存在多稳态,并确定其稳态类型;2)通过监测潮间带生态系统水动力过程、沉积动力过程以及盐沼植物生长和扩散情况,分析盐沼植被与沉积地貌之间的正反馈作用,进而探讨潮间带生态系统多稳态的形成机制.结果表明: 1)潮间带生态系统的归一化植被指数(NDVI)频度分布存在明显的双峰特征,且盐沼植物成活存在生物量阈值效应,均证实潮间带生态系统存在多稳态,“盐沼”和“光滩”是潮间带生态系统的两种相对稳定状态;2)崇明东滩盐沼前沿的沉积地貌表现出泥沙快速淤积的趋势,显著促进了盐沼植物的生长,盐沼植物与泥沙淤积之间的这种正反馈作用是潮间带生态系统形成多稳态的主要原因;3)盐沼植被扩散格局监测结果在景观尺度上也表明,泥沙淤积作用促进了潮间带生态系统“盐沼”和“光滩”多稳态的形成.本研究既丰富了滨海湿地稳态转换的机理研究,也为我国开展海岸带保护、修复和管理提供了科学依据,具有重要的理论和实践意义.     

Ecosystem spatial self-organization: Free order for nothing?

Ecosystems are complex adaptive systems (CAS) by nature, which means that macroscopic patterns and properties emerge from, and feed back to affect, the interactions among adaptive individual ecological agents. These agents then further adapt (genetically) to the outcomes of those interactions. The concept of self-organization has become increasingly important for understanding ecosystem spatial heterogeneity and its consequences. It is well accepted that ecosystems can self-organize, and that resulting spatial structures carry functional consequences. Feedbacks from the outcome of spatial pattern to the individual agents from which patterns emerge, are an essential component of the definition of CAS but have been rarely examined for ecosystems. We explore whether spatial self-organization provides a mechanism for such feedback for ecosystems as CAS, that is, whether ecosystem-level outcomes of self-organized patterning could feed back to affect or even reinforce local pattern-forming processes at the agent level. Diffuse feedbacks of ecological and evolutionary significance ensue as a result of spatial heterogeneity and regular patterning, whether this spatial heterogeneity results from an underlying template effect or from self-organization. However, feedbacks directed specifically at pattern-forming agents to enhance pattern formation—reinforcing feedback—depend upon the level of organization of agents. Reinforcing evolutionary feedbacks occur at the individual level or below. At the ecosystem level, evidence for mechanisms of feedback from outcomes to patterning to agents forming the patterning remain tenuous. Spatial self-organization is a powerful dynamic in ecosystem and landscape science but feedbacks have been only loosely integrated so far. Self-organized patterns influencing dynamics at the ecosystem level represent “order for free”. Whether or not this free order generated at the ecosystem level carries evolutionary function or is merely epiphenomenal is a fundamental question that we address here.     

火在生态系统中的作用

前言火对植被的影响是在近十多年来才被完全确认的。在二十世纪的大部分时间内,火常常被认为是破坏生态系统的非自然因子;然而,植被学家、人类学家、地理学家和其他许多学者都认为北美的土族印地安人和其他许多民族曾广泛地利用火作为土地管理的工具(Van-     

Microbial carbon limitation: The need for integrating microorganisms into our understanding of ecosystem carbon cycling

Numerous studies have demonstrated that fertilization with nutrients such as nitrogen, phosphorus, and potassium increases plant productivity in both natural and managed ecosystems, demonstrating that primary productivity is nutrient limited in most terrestrial ecosystems. In contrast, it has been demonstrated that heterotrophic microbial communities in soil are primarily limited by organic carbon or energy. While this concept of contrasting limitations, that is, microbial carbon and plant nutrient limitation, is based on strong evidence that we review in this paper, it is often ignored in discussions of ecosystem response to global environment changes. The plant‐centric perspective has equated plant nutrient limitations with those of whole ecosystems, thereby ignoring the important role of the heterotrophs responsible for soil decomposition in driving ecosystem carbon storage. To truly integrate carbon and nutrient cycles in ecosystem science, we must account for the fact that while plant productivity may be nutrient limited, the secondary productivity by heterotrophic communities is inherently carbon limited. Ecosystem carbon cycling integrates the independent physiological responses of its individual components, as well as tightly coupled exchanges between autotrophs and heterotrophs. To the extent that the interacting autotrophic and heterotrophic processes are controlled by organisms that are limited by nutrient versus carbon accessibility, respectively, we propose that ecosystems by definition cannot be ‘limited’ by nutrients or carbon alone. Here, we outline how models aimed at predicting non‐steady state ecosystem responses over time can benefit from dissecting ecosystems into the organismal components and their inherent limitations to better represent plant–microbe interactions in coupled carbon and nutrient models.     

Moving from autonomous to planned adaptation in the montane forests of southeastern Australia under changing fire regimes

Forest ecosystems and their associated natural, cultural and economic values are highly vulnerable to climate driven changes in fire regimes. A detailed knowledge of forest ecosystem responses to altered fire regimes is a necessary underpinning to inform options for adaptive responses under climate change, as well as for providing a basis for understanding how patterns of distribution of vegetation communities that comprise montane forest ecosystems may change in the future. Unplanned consequential adaptation of both natural and human systems, i.e. autonomous adaptation, will occur without planned intervention, with potentially negative impacts on ecosystem services. The persistence of forest stands under changing fire regimes and the maintenance of the ecosystem services that they provide pivot upon underlying response traits, such as the ability to resprout, that determine the degree to which composition, structure and function are likely to change. The integration of ecosystem dynamics into conceptual models and their use in exploring adaptation pathways provides options for policy makers and managers to move from autonomous to planned adaptation responses. Understanding where autonomous adaptation provides a benefit and where it proves potentially undesirable is essential to inform adaptation choices. Plausible scenarios of ecological change can be developed to improve an understanding of the nature and timing of interventions and their consequences, well before natural and human systems autonomously adapt in ways that may be detrimental to the long‐term provision of ecosystem services. We explore the utility of this approach using examples from temperate montane forest ecosystems of southeastern Australia.     

Gaia as a complex adaptive system

We define the Gaia system of life and its environment on Earth, review the status of the Gaia theory, introduce potentially relevant concepts from complexity theory, then try to apply them to Gaia. We consider whether Gaia is a complex adaptive system (CAS) in terms of its behaviour and suggest that the system is self-organizing but does not reside in a critical state. Gaia has supported abundant life for most of the last 3.8 Gyr. Large perturbations have occasionally suppressed life but the system has always recovered without losing the capacity for large-scale free energy capture and recycling of essential elements. To illustrate how complexity theory can help us understand the emergence of planetary-scale order, we present a simple cellular automata (CA) model of the imaginary planet Daisyworld. This exhibits emergent self-regulation as a consequence of feedback coupling between life and its environment. Local spatial interaction, which was absent from the original model, can destabilize the system by generating bifurcation regimes. Variation and natural selection tend to remove this instability. With mutation in the model system, it exhibits self-organizing adaptive behaviour in its response to forcing. We close by suggesting how artificial life ('Alife') techniques may enable more comprehensive feasibility tests of Gaia.     

Tipping point in plant–fungal interactions under severe drought causes abrupt rise in peatland ecosystem respiration

Ecosystems are increasingly prone to climate extremes, such as drought, with long‐lasting effects on both plant and soil communities and, subsequently, on carbon (C) cycling. However, recent studies underlined the strong variability in ecosystem's response to droughts, raising the issue of nonlinear responses in plant and soil communities. The conundrum is what causes ecosystems to shift in response to drought. Here, we investigated the response of plant and soil fungi to drought of different intensities using a water table gradient in peatlands—a major C sink ecosystem. Using moving window structural equation models, we show that substantial changes in ecosystem respiration, plant and soil fungal communities occurred when the water level fell below a tipping point of ?24 cm. As a corollary, ecosystem respiration was the greatest when graminoids and saprotrophic fungi became prevalent as a response to the extreme drought. Graminoids indirectly influenced fungal functional composition and soil enzyme activities through their direct effect on dissolved organic matter quality, while saprotrophic fungi directly influenced soil enzyme activities. In turn, increasing enzyme activities promoted ecosystem respiration. We show that functional transitions in ecosystem respiration critically depend on the degree of response of graminoids and saprotrophic fungi to drought. Our results represent a major advance in understanding the nonlinear nature of ecosystem properties to drought and pave the way towards a truly mechanistic understanding of the effects of drought on ecosystem processes.     

Molecular ecosystems

Biologists employ a suggestive metaphor to describe the complexities of molecular interactions within cells and embryos: cytological components are said to be part of “ecosystems” that integrate them in a complex network of relations with many other entities. The aim of this essay is to scrutinize the molecular ecosystem, a metaphor that, despite its longstanding history, has seldom be articulated in detail. I begin by analyzing some relevant analogies between the cellular environment and the biosphere. Next, I discuss the applicability of the molecular ecosystem concept in actual scientific practice.     

The ephemeral resource patch concept

Ephemeral resource patches (ERPs) – short lived resources including dung, carrion, temporary pools, rotting vegetation, decaying wood, and fungi – are found throughout every ecosystem. Their short-lived dynamics greatly enhance ecosystem heterogeneity and have shaped the evolutionary trajectories of a wide range of organisms – from bacteria to insects and amphibians. Despite this, there has been no attempt to distinguish ERPs clearly from other resource types, to identify their shared spatiotemporal characteristics, or to articulate their broad ecological and evolutionary influences on biotic communities. Here, we define ERPs as any distinct consumable resources which (i) are homogeneous (genetically, chemically, or structurally) relative to the surrounding matrix, (ii) host a discrete multitrophic community consisting of species that cannot replicate solely in any of the surrounding matrix, and (iii) cannot maintain a balance between depletion and renewal, which in turn, prevents multiple generations of consumers/users or reaching a community equilibrium. We outline the wide range of ERPs that fit these criteria, propose 12 spatiotemporal characteristics along which ERPs can vary, and synthesise a large body of literature that relates ERP dynamics to ecological and evolutionary theory. We draw this knowledge together and present a new unifying conceptual framework that incorporates how ERPs have shaped the adaptive trajectories of organisms, the structure of ecosystems, and how they can be integrated into biodiversity management and conservation. Future research should focus on how inter- and intra-resource variation occurs in nature – with a particular focus on resource × environment × genotype interactions. This will likely reveal novel adaptive strategies, aid the development of new eco-evolutionary theory, and greatly improve our understanding of the form and function of organisms and ecosystems.     

Revisiting the ‘bank of microscopic forms’ in macroalgal‐dominated ecosystems

Theoretical ecological models, such as succession and facilitation, were defined in terrestrial habitats, and subsequently applied to marine and freshwater habitats in intertidal and then subtidal realms. One such model is the soil seed bank, defined as all viable seeds (or fruits) found near the soil surface that facilitate community restoration/recovery. “Banks of microscopic forms” have been hypothesized in aquatic habitats and recent work from aquaculture has highlighted dormancy in algal life cycle stages. To reinvigorate the discussions about these algal banks, we discuss differences in life cycles, dispersal, and summarize research on banks of macroalgal stages in aquatic ecosystems that may be easier to explore with modern advances in molecular technology. With focus on seminal work in global kelp forest ecosystems, we present a pilot study in northern California as proof of concept that Nereocystis luetkeana and Alaria marginata stages can be detected within kelp forests in the biofilm of rocks and bedrock using targeted primers long after zoospore release. Considering the increased interest in algae as an economic resource, [blue] carbon sink, and as ecosystem engineers, the potential for “banking” macroalgal forms could be a mechanism of resilience and recovery in aquatic populations that have complex life cycles and environmental cues for reproduction. Molecular barcoding is becoming an important tool for identifying banks of macroalgal forms in marine communities. Understanding banks of macroalgal stages, especially in deforested habitats with intense disturbance and grazer pressure, will allow researchers and marine resource managers to facilitate this natural process in recovery of the aquatic system.     

生态系统的多稳态与突变

许多生态系统可能在短时间内发生难以预料的状态突变, 其中一些生态系统突变的机理可以用多稳态理论进行解释。近年来生态系统的多稳态和突变现象及其机理吸引了研究者和管理者的广泛关注。本文重点对生态系统多稳态的理论基础、识别方法及稳态转换发生的早期预警信号进行综述, 并基于典型生态系统过程对现实世界中可能观测到的稳态转换进行实例分析, 最后对多稳态概念框架和理论应用中的潜在争议进行讨论, 以期为非线性生态系统动态的理论研究、管理实践和生物多样性保护等提供参考。     

Landscape‐scale determinants of non‐native fish communities

Aim Predicting and preventing invasions depends on knowledge of the factors that make ecosystems susceptible to invasion. Current studies generally rely on non‐native species richness (NNSR) as the sole measure of ecosystem invasibility; however, species identity is a critical consideration, given that different ecosystems may have environmental characteristics suitable to different species. Our aim was to examine whether non‐native freshwater fish community composition was related to ecosystem characteristics at the landscape scale. Location United States. Methods We described spatial patterns in non‐native freshwater fish communities among watersheds in the Mid‐Atlantic region of the United States based on records of establishment in the U.S. Geological Survey’s Nonindigenous Aquatic Species Database. We described general relationships between non‐native species and ecosystem characteristics using canonical correspondence analysis. We clustered watersheds by non‐native fish community and described differences among clusters using indicator species analysis. We then assessed whether non‐native communities could be predicted from ecosystem characteristics using random forest analysis and predicted non‐native communities for uninvaded watersheds. We estimated which ecosystem characteristics were most important for predicting non‐native communities using conditional inference trees. Results We identified four non‐native fish communities, each with distinct indicator species. Non‐native communities were predicted based on ecosystem characteristics with an accuracy of 80.6%, with temperature as the most important variable. Relatively uninvaded watersheds were predicted to be invasible by the most diverse non‐native community. Main conclusions Non‐native species identity is an important consideration when assessing ecosystem invasibility. NNSR alone is an insufficient measure of invasibility because ecosystems with equal NNSR may not be equally invasible by the same species. Our findings can help improve predictions of future invasions and focus management and policy decisions on particular species in highly invasible ecosystems.     

A review of important concepts in the trophic organization of pelagic ecosystems

The physical environment has an important influence on the size composition of primary producers in plankton communities. This effect is transmitted through the trophic structure by size selective feeding processes at each level. A pictorial representation of trophic organization in pelagic ecosystems is presented which illustrates the complex involvement of individual species in the food web and the dynamic nature of ecosystem development along alternate pathways.This research was supported in full by the National Science Foundation, Grant No. NSF OCE76-02035.