Effectiveness of entropy-based functions in the analysis of ecosystem state and development

Following the advances in the field of the thermodynamics of far-from-equilibrium systems, several ecological orientors (indicators able to describe the stage and orientation of ecosystem development) incorporating entropy terms have been proposed. Although most of the proposed functions have a good theoretical basis and have proved to perform adequately as ecological indicators, their suitability as ecological orientors has yet to receive a full confirmation in real case studies. The aim of the present contribution is to examine how several entropy-based indicators (exergy, structural information, entropy production, specific entropy production and the Eco-exergy index) perform as orientors when applied to a special case of ecological succession, i.e. eutrophication, in a homogeneous set of shallow lakes lying along a trophic gradient, from oligotrophy to hyper-eutrophy. The results show that a coherent pattern of response emerge, which is also consistent with the classical ecological theory. In particular, the maximisation of storage and the minimisation of specific entropy production are confirmed as the most reliable principles of ecosystem development, whereas the maximisation of dissipation (as entropy production) appears as a debatable criterion of development.     

Complementarity of ecological goal functions

This paper summarizes, in the framework of network environ analysis, a set of analyses of energy-matter flow and storage in steady-state systems. The network perspective is used to codify and unify ten ecological orientors or extremal principles: maximum power (Lotka), maximum storage (J?rgensen-Mejer), maximum empower and emergy (Odum), maximum ascendency (Ulanowicz), maximum dissipation (Schneider-Kay), maximum cycling (Morowitz), maximum residence time (Cheslak-Lamarra), minimum specific dissipation (Onsager, Prigogine), and minimum empower to exergy ratio (Bastianoni-Marchettini). We show that, seen in this framework, these seemingly disparate extrema are all mutually consistent, suggesting a common pattern for ecosystem development. This pattern unfolds in the network organization of systems.     

The strategy of ecosystem development: specific dissipation as an indicator of ecosystem maturity

The ratio of entropy generation rate to entropy embodied in structures relatively to the surroundings can be considered as an indicator of the ability of a self-organizing dissipative system to maintain itself far from equilibrium by pumping out entropy. The higher the ratio (which may be called the specific entropy production or the specific dissipation of a system), the lower the capacity of a system to convert the incoming low-entropy energy into internal organization. It appears that the ratio attains special significance for interpreting the evolution of biological systems, as the maximum expression of self-organizing systems, from the sub-cellular to the ecosystem scale. This paper proposes specific dissipation, written as the ratio of biological entropy production to exergy stored in the living biomass, as a thermodynamic orientor as well as an indicator of the development state of ecological systems. After having presented a method for estimating the specific dissipation in lakes, the adequacy of the proposed indicator is discussed and also tested by comparing its response to those of some classical ecological attributes (successional sequences of species, biodiversity, individual body size, structural organization and generation time of organisms) throughout the seasonal progression of the plankton community in Lake Trasimeno (Umbria, Italy). The results support the hypothesis that the minimization of specific dissipation is a primary criterion of evolution of ecological systems and also sustain the use of specific dissipation as an indicator of ecological maturity.     

Does the intermediate disturbance hypothesis comply with thermodynamics?

A unique data set from Keszthely Bay, Lake Balaton has been applied to develop a dynamic structural model able to describe the observed changes in phytoplankton biomass and diversity. We tested whether the model reacts according to the Intermediate Disturbance Hypothesis and according to the hypothesis that ecosystem reactions attempt to maximize the thermodynamic function exergy under prevailing conditions. If the answer to these tests are confirmatory, it can be considered a support for IDH and for the use of the exergy maximization principle as a general principle to explain ecosystem reactions.     

森林生态系统发展和植物种群变化的热力学过程

随着生态学的发展,人们对自然生态系统的认识逐渐从对自然现象的记录、描述,发展为对机制的系统认识。热力学定律为人们提供了认识系统发展规律的理论基础,但在生态系统中的应用还处于起步阶段。基于前人关于生态系统可用能的研究。探讨了森林生态系统和植物种群变化的热力学过程。在生态系统水平上,把可用能耗散分为了植物耗散和环境耗散两个部分,并探讨了这两个过程之间的关系。第一次明确地提出蒸散是植物耗散的主要部分。在植物种群水平上,“可用能储存”与“可用能耗散”是决定植物竞争力的关键因子,在同一区域相同条件下,拥有更大可用能耗散能力的物种应当被优先选择。因此,群落中的优势物种应当比同层次的伴生种具有相对高的生长速度和更强的蒸腾能力。研究试图在热力学理论体系与实际生态系统的生理生态过程之间建立了纽带和桥梁,为开展森林生态系统的健康评估、深刻认识植物与环境的关系、以及评价物种竞争能力提供新的理论视野。     

Ecological complexity effects on thermal signature of different Madeira island ecosystems

From a systemic perspective, evolution and natural succession promote the creation of efficient biological structures and processes that capture and dissipate the solar energy, maximizing the entropy production. This ecological complexification results in better ecosystem thermodynamic performance indicated by lower temperature.In a brief period of evolutionary time human-induced disturbance has altered profoundly the structure and functioning of the Earth System, i.e. ecological simplification.The objective is to understand whether remote sensing data can be considered appropriate proxy indicators to test if more mature and complex ecosystems have higher entropy production rates which lead to lower and attenuated ecosystem temperatures.Simple remote sensing measurements of Madeira Island for Thermal Infrared Radiation and Normalized Difference Vegetation Index were used to analyse the surface temperature and biomass cover of Madeira ecosystems spectrum of different states of human-induced disturbance.The findings revealed it was possible to distinguish between ecosystem types using thermodynamic indicators, where older ecosystems with more complex structures exhibit more attenuated lower average temperatures.It was also found that habitat heterogeneity can represent either artificial (human) or natural disturbance with opposite consequences in the ecosystem thermal signature, i.e. lower temperature when natural disturbance and higher if anthropogenic disturbance.Simple thermal remote sensing data can be used as systemic indicator of ecosystem health by reflecting it levels of eco-exergy, i.e the available work energy in the ecosystem.     

Ecological succession as an energy dispersal process

Ecological succession is described by the 2nd law of thermodynamics. According to the universal law of the maximal energy dispersal, an ecosystem evolves toward a stationary state in its surroundings by consuming free energy via diverse mechanisms. Species are the mechanisms that conduct energy down along gradients between repositories of energy which consist of populations at various thermodynamic levels. The salient characteristics of succession, growing biomass production, increasing species richness and shifting distributions of species are found as consequences of the universal quest to diminish energy density differences in least time. The analysis reveals that during succession the ecosystem's energy transduction network, i.e., the food web organizes increasingly more effective in the free energy reduction by acquiring new, more effective and abandoning old, less effective species of energy transduction. The number of species does not necessarily peak at the climax state that corresponds to the maximum-entropy partition of species maximizing consumption of free energy. According to the theory of evolution by natural selection founded on statistical physics of open systems, ecological succession is one among many other evolutionary processes.     

Total energy costs in ecosystems

A micro and macro ecological system hypotheses and their associated theory of total (direct and indirect) energy cost have been proposed to explain the meaning of system growth and the steady state condition. Models have been proposed which can be used to test the hypotheses. In brief, the hypotheses are that while the components of an ecosystem strive to maximize their total direct and indirect energy storage within the constraints of their production characteristics, the overall system strives to minimize the metabolized energy per unit of stored biomass energy. The steady state result is a system with the largest possible stored biomass under the general constraints of water, air, soil and sunlight availability.Feeding strategies are discussed with the conclusion that a net total energy-maximizing feeding strategy may be able to replace the present concepts.     

Successional changes in the morphology and ecological responses of a grazed pasture ecosystem in Central Spain

Morphological characteristics related to spatial occupation, reproduction and adaptations to grazing were used to characterize the most frequent species in a therophytic pastureland of Central Spain.Periodic ploughing is a traditional practice in these pastures and allows observation of successional change. In the present study, four neighbouring slopes of differing time since last ploughing were chosen. Species biomass was measured at different times during the annual growing season for two different slope positions.Grazing pressure is an important environmental factor affecting ecosystem organization, the most palatable plants tending to show increasing biomass with succession. In the most mature stages, there is a predominance of species characterized by horizontal occupation of space and sprouting after mowing or grazing.During succession segregation of the different morphological characteristics occurs in slope sectors related to geomorphological dynamics. Similarly, phenological development tends to be later in pastures in the lowest slope zones, due probably to their greater summer soil moisture content.Nomenclature follows: Tutin et al., 1964–1980. Flora Europaea.     

水稻-褐飞虱-八斑球腹蛛生态系统的能流

为了使水稻害虫综合治理提高到系统工程的水平,作者在室内30℃恒温下,测定了褐飞虱和八斑球腹蛛各发育阶段的虫(蛛)体组织潜能量、生殖生产量、呼吸量、蜕及蛛丝含能量。根据晚稻田褐飞虱和八斑球腹蛛种群数量的系统调查结果,以及水稻生物量与热值的测定值,给出了水稻-褐飞虱-八斑球腹蛛生态系统的能量流动图,分析了该生态系统的净生态效率、对光能或潜能的利用效率和生物量、能量生态锥体。     

Does microbial life always feed on negative entropy? Thermodynamic analysis of microbial growth.

Schr?dinger stated in his landmark book, What is Life?, that life feeds on negative entropy. In this contribution, the validity of this statement is discussed through a careful thermodynamic analysis of microbial growth processes. In principle, both feeding on negative entropy, i.e. yielding products of higher entropy than the substrates, and generating heat can be used by microorganisms to rid themselves of internal entropy production resulting from maintenance and growth processes. Literature data are reviewed in order to compare these two mechanisms. It is shown that entropy-neutral, entropy-driven, and entropy-retarded growth exist. The analysis of some particularly interesting microorganisms shows that enthalpy-retarded microbial growth may also exist, which would signify a net uptake of heat during growth. However, the existence of endothermic life has never been demonstrated in a calorimeter. The internal entropy production in live cells also reflects itself in the Gibbs energy dissipation accompanying growth, which is related quantitatively to the biomass yield. An empirical correlation of the Gibbs energy dissipation in terms of the physico-chemical nature of the growth substrate has been proposed in the literature and can be used to predict the biomass yield approximately. The ratio of enthalpy change and Gibbs energy change can also be predicted since it is shown to be approximately equal to the same ratio of the relevant catabolic process alone.     

生态能质(eco-exergy)在水生生态系统建模和评价中的应用

生态能质(eco-exergy)是指系统从给定状态达到热力学平衡状态所做的功,可以度量生态系统的复杂生物化学组分及生态结构。系统具有的能质越大,有序化程度越高,稳定性也越强。生态能质和比生态能质(specific eco-exergy)指标能够评价水生生态系统健康状况,对水生生态系统演替阶段具有指示作用。本文阐述了生态能质的定义、生态学意义以及生态能质值和权重因子fi值的计算方法。对生态能质作为功能函数在水生生态系统结构动力学模型(structural dynamic models,SDMs)中以及作为生态指标在生态系统健康评价、生态恢复研究中的实际应用进行了总结。最后探讨了生态能质指标的局限性以及未来的发展。此外,文中建议将eco-exergy和specific eco-exergy统一翻译为生态能质和比生态能质。     

Thermodynamic stability of ecosystems

The stability of ecosystems during periods of stasis in their macro-evolutionary trajectory is studied from a non-equilibrium thermodynamic perspective. Individuals of the species are considered as units of entropy production and entropy exchange in an open thermodynamic system. Within the framework of the classical theory of irreversible thermodynamics, and under the condition of constant external constraints, such a system will naturally evolve toward a globally stable thermodynamic stationary state. It is thus suggested that the ecological steady state, or stasis, is a particular case of the thermodynamic stationary state, and that the evolution of community stability through natural selection is a manifestation of non-equilibrium thermodynamic directives. Furthermore, it is argued that punctuation of stasis leading to ecosystem succession, may be a manifestation of non-equilibrium "phase transitions" brought on by a change of external constraints through a thermodynamic critical point.     

Soil fertility effects on tree seedling performance are light‐dependent: evidence from a long‐term soil chronosequence

Soil chronosequences are a powerful tool for understanding how limitation of plant growth by nutrients and light changes throughout ecosystem development, but experimental tests of how availability of these resources interact to influence plant performance as ecosystem development proceeds are rare. We utilise the well‐characterised Franz Josef soil chrononosequence in New Zealand, a sequence of sites caused by a retreating glacier that spans ca 120 000 years and that includes all stages of ecosystem development from primary succession through to retrogression. Soil fertility is relatively low at either end of the sequence due to limitation of biological processes initially by N and ultimately by P whereas light availability is lowest at intermediate stages of the sequence dominated by tall forest. Growth and leaf traits of nine woody plant species, including those that occur widely along the chronosequence and those that are restricted to short portions of it, were quantified in a mesocosm experiment. Phytometers of these species were each grown in each of nine soils collected from throughout the chronosequence at either high (30%) or low (2%) light levels; these soil and light conditions represent the full variation observed along the sequence. Plant growth and biomass were greatest in soils from intermediate stages of the chronosequence and in high light. However, the stimulatory effects of soil fertility largely disappeared under shaded conditions that are characteristic of intermediate stages of ecosystem development. Our results demonstrate that long‐term changes in soil fertility and light availability that occur throughout ecosystem development had direct effects on plant species performance, but that there were stronger interactive effects of soils and light availability. Because light and soil resource availability shift predictably but have different trajectories throughout ecosystem development, our results help to understand variation in plant species performance and community assembly along complex environmental gradients.     

青海海北地区矮嵩草草甸生物量和能量的分配

 此项研究工作于1980年在海北高寒草甸生态系统定位站进行。本文研究了青藏高原地区分布面积广、草质优良,在畜牧业生产中有重要意义的矮嵩草草甸的生物量和它的能量分配关系,测定了地上,地下生物量和不同物候期主要植物类群的热值含量。研究结果表明：矮嵩草草甸生物量的季节动态较为明显,地上生物量随生长季节的水热条件和植物的生长发育阶段而变化,9月初地上生物量达到峰值(296.66g／m2),此后生物量逐渐减少,到枯黄前而停止;地下根系生物量在返青期较高,生长旺盛期最低,枯黄期最高,这同植物生长发育阶段的物质运转有关。矮嵩草草甸主要植物类群的热值以生长旺盛期最高,枯黄期次之,返青期较低;各类草的热值,以莎草类最高,禾草类次之,杂类草最低。矮嵩草草甸总初级生产量为909.49g／m2·年,其中地上为296.66g／m2·年,地下为596.67g／m2·年,枯枝落叶为16.16g／m2·年。群落在不同生长期所固定的太阳能数值不一,以枯黄前所固定的太阳能为最多,生长期整个群落的光能利用率为0.295%。     

Leaf-nutrient accumulation and turnover at three stages of succession from heathland to forest

Abstract. Accumulation of nutrients in leaves of the dominating species of three ecosystems, characterizing the secondary succession from Genisto-Callunetum heathland through Leucobryo-Pinetum birch-pine woodland to mature Querco-Fagetum oak-beech forest, as well as nutrient turnover within these ecosystems was studied. The objective of the study was to establish potential variations in quantity and quality of nutrient supply to the plants with respect to succession dynamics. The results show very low leaf nutrient concentrations of all species investigated, coinciding with low nutrient availability in the soil. However, the nutrient content of leaves and leaf litter of Quercus petraea and Fagus sylvatica, which dominate the late succession stages, and in Betulapéndula are higher than in the photosynthetic organs (leaves and young shoots) of Calluna vulgaris and Pinus sylvestris. The combination of the higher nutrient content of the leaves and an increasing leaf-litter production during succession results in an increased nutrient turnover via leaf-litter fall. However, due to the high leaf biomass, the storage of nutrients in the leaf biomass is highest within the birch-pine woodland. From this, it may be assumed that the low demand and the low loss of nutrients via leaf-litter fall are favourable for Pinus at the early stages of forest succession on poor sandy soils. In contrast, Quercus and Fagus are provided with better growth conditions at the later stages of succession resulting from the accumulation of plant-available nutrients in the ecosystem by Pinus sylvestris, combined with a higher nutrient turnover as compared with the heathland.     

Stability of foodwebs and its relation to species diversity

The stability of the ecosystem is defined in terms of its ensemble of essentially similar states. This definition, along with the conservation laws of matter and energy, is used to classify the possible mechanisms of ecological stability. These include the development of balance, physiological adaptability (apropos variability in the conditions affecting the use of energy as well as variability in available energy), selective distribution of community fluctuation, and multifunctionality. In particular selective distribution of fluctuation can be achieved through the development of threshold behavior, i.e. sacrifice of energy to detritus pathways. Overlapping multifunctionality coupled with this threshold behavior allows control over the possible pathways of energy flow. The threshold behavior is based on intra- or interspecies communication. The development of these mechanisms in the course of ecological succession is examined under the assumption that the time average of the energy content of the ecosystem tends to a maximum. This development has implications as regards the relative importance of detritus pathways and the number of grazing pathway species during various stages of succession.     

森林次生演替和土壤层次对微生物群落结构的影响

森林次生演替与生态系统结构和功能的动态变化密切相关。大多数研究主要关注植物群落以及土壤有机碳(SOC)的变化,然而土壤微生物群落如何响应森林次生演替还需要进一步探究。本研究以长白山森林次生演替序列(20、80、120、200和≥300年)和两个土壤层次为对象,采用磷脂脂肪酸微生物标志物,探究温带森林次生演替过程中地下微生物群落结构变化。森林次生演替改变了土壤微生物群落结构,主要归因于某些特定微生物类群的变化,演替前期革兰氏阴性菌和腐生真菌占主导,而在演替后期革兰氏阳性菌和丛枝菌根真菌占主导。另外,土壤有机质数量和质量差异是影响微生物群落结构和生物量的主要环境因素。森林演替前期和中期增加的SOC含量促进了微生物生物量,而演替后期增加的难分解芳香族有机组分抑制了微生物生物量合成。土壤层次间理化性质的差异导致微生物群落变化,有机质层高的SOC以及氮含量导致更多微生物生物量的合成。微生物群落在时间和空间尺度的变化及其驱动因素反映了生态系统结构和功能对环境变化的响应。     

人类活动对青藏高原高寒矮嵩草草甸碳过程的影响

随着人类活动干扰(放牧)的增加,青藏高原高寒嵩草甸的退化演替过程依次为禾草-矮嵩草群落、矮嵩草群落、小嵩草群落和杂类草-黑土滩4个阶。其中小嵩草群落又可划分为草毡表层加厚、开裂与塌陷3个子阶段。采用时空转换的方法,研究了人类活动对青藏高原高寒矮嵩草草甸碳过程的影响。结果表明,随着人类干扰强度的增加,植物群落地上部分有机碳储量逐渐降低,由禾草-矮嵩草群落的(134.7±17.3)gC/m2逐渐降低到杂类草-黑土滩次生裸地(18.96±6.18)gC·m-2。土壤、植物地下部分有机碳贮量呈单峰曲线变化,草毡表层开裂子阶段最高,分别为(49.7±0.83)gC·kg-1和(3596.7±179.8)gC·m-2。;杂类草-黑土滩阶段最低,分别为(19.2±1.13)gC·kg-1和(121.6±6.1)gC·m-2。受植物地下部贮碳的影响,土壤-植被系统呈现逐渐降低的变化特征。随人类活动干扰的加强,高寒嵩草草地植物有机碳地下/地上分配比发生巨大改变,草地草毡表层厚度不高于4.3cm是保证草地生产与生态服功能双赢的重要指标。     

Oomycetes along a 120,000 year temperate rainforest ecosystem development chronosequence

The occurrence of plant-associated oomycetes in natural ecosystems and particularly during long-term ecosystem development is largely unknown. Using DNA sequencing, we investigated the frequency and host relationships of plant-root-associated oomycete communities along a 120 000 y glacial chronosequence, comprising site ages with rapid compositional change (“early succession”; 5–70 y old soil); relatively stable higher-diversity sites (“mature”, 280–12000 y); and ancient, nutrient-limited soils with declining plant biomass and stature (“retrogression”, 60 000, 120 000 y). Plant-associated oomycetes were frequent in early successional sites, occurring in 38–65% of plant roots, but rare (mean 3%) in all older ecosystems. Oomycete OTUs occurred non-randomly with plant host species, and were more frequent on those plant species that declined most strongly in abundance between ecosystem ages. While oomycetes were common in early succession, their absence in older sites suggests a limited role in later stages of ecosystem development.