土壤动物粒径谱研究进展

群落结构如何响应环境变化是生态学研究长期关注的核心问题之一。粒径谱由个体大小和多度构建而来,与营养级转换速率相关、反映生态系统过程动态以及表征生态系统稳定性,可以将其视为一个综合的功能多样性指标用于预测和表征群落的组成以及生态系统功能如何响应环境压力。粒径谱研究最初始于水生生态系统,近年来被引入到土壤动物群落生态学的研究中。简要回顾粒径谱的概念由来及理论基础,分析比较了当前粒径谱研究中的4种易混淆类型,介绍了常用的两类土壤动物粒径谱构建方法及其生态学意义,梳理了土壤动物粒径谱对环境梯度响应与生态化学计量学相结合的研究进展,并指出了应用粒径谱研究土壤动物群落的难点及限制条件。未来,在基础理论研究方面,土壤动物粒径谱应关注个体大小与营养级位置及能量利用关系;在应用方面,土壤动物粒径谱可结合传统的分类方法广泛应用于指示环境污染、生态恢复、保育生物以及土地利用变化等。     

Modelling the effects of climate change on the distribution and production of marine fishes: accounting for trophic interactions in a dynamic bioclimate envelope model

Climate change has already altered the distribution of marine fishes. Future predictions of fish distributions and catches based on bioclimate envelope models are available, but to date they have not considered interspecific interactions. We address this by combining the species‐based Dynamic Bioclimate Envelope Model (DBEM) with a size‐based trophic model. The new approach provides spatially and temporally resolved predictions of changes in species' size, abundance and catch potential that account for the effects of ecological interactions. Predicted latitudinal shifts are, on average, reduced by 20% when species interactions are incorporated, compared to DBEM predictions, with pelagic species showing the greatest reductions. Goodness‐of‐fit of biomass data from fish stock assessments in the North Atlantic between 1991 and 2003 is improved slightly by including species interactions. The differences between predictions from the two models may be relatively modest because, at the North Atlantic basin scale, (i) predators and competitors may respond to climate change together; (ii) existing parameterization of the DBEM might implicitly incorporate trophic interactions; and/or (iii) trophic interactions might not be the main driver of responses to climate. Future analyses using ecologically explicit models and data will improve understanding of the effects of inter‐specific interactions on responses to climate change, and better inform managers about plausible ecological and fishery consequences of a changing environment.     

湛江高桥红树林湿地底栖动物粒径谱

在我国,红树林湿地底栖动物粒径谱研究很少。根据2010年1月、4月、7月、10月在湛江高桥红树林湿地获得的大型和小型底栖动物数据,构建了底栖动物生物量粒径谱,以期为湛江高桥红树林湿地的生态保护和持续利用提供科学依据。主要研究结果如下:(1)高桥红树林湿地生物量粒径谱基本为3峰模式。第一峰在-2粒级,主要由线虫构成;第二峰在4—12粒级,主要由寡毛类、多毛类和小个体甲壳类构成;第三峰在13—22粒级,主要由大个体腹足类、双壳类和甲壳类构成。(2)木榄、桐花树和无瓣海桑生境在0—4粒级之间出现一个明显的波谷,这个波谷介于线虫和寡毛类之间,是大型与小型底栖动物粒级交汇区。(3)高桥红树林湿地底栖动物正态化生物量粒径谱的斜率大于-1,截距为16.533—18.150。桐花树(Aegiceras corniculatum)和无瓣海桑(Sonneratia apetala)生境的截距、最小粒级的生物量(BMS)高于木榄(Bruguiera gymnorrhiza)和盐地鼠尾粟(Sporobolus virginicus)生境,说明桐花树和无瓣海桑生境的底栖动物生产力水平较木榄和盐地鼠尾粟生境的高;秋季的截距、BMS较其他季节高,说明秋季的底栖动物生产力水平较其他季节高。     

Biomass size spectra of net plankton in the adjacent area near the Yangtze River Estuary in spring

Based on data from survey carried out in spring 2005, the biomass size spectra of net plankton were explored in the adjacent sea of Yangtze River Estuary. Results revealed an approximately continuous size distribution of plankton individuals, from phytoplankton (5–250 μm cell^?1 in equivalent sphere diameter (ESD), 15 pg–146 ng C cell^?1) to zooplankton (120 μm–2 cm ESD, 115 ng–7.5 mg C ind^?1). The normalized spectra (carbon scale) were linear with slope ranged from –0.889 to –0.445, and intercept ranged from 12.866 to 16.863 (all stations together, slope = –0.606, intercept = 19.448), indicating strong deviations from the ideal value (slope = –1.22) of a steady pelagic ecosystem. Correlation analysis presented that intercept and regression coefficient of net plankton size spectra had significant relationship with plankton biomass size diversity.     

Three into two doesn't go: two-dimensional models of bird eggs, snail shells and plant roots

Three published two-dimensional analyses of three-dimensional structures are reinvestigated. (1) In their 1997 paper, Barta & Szekely sought shapes of bird eggs that maximized the size of eggs that packed under a circular brood patch. Inappropriately they measured egg size by cross-sectional area; maximizing volume implies different optima. Also their genetic algoridim mislocated their optima, probably because of too much mutation. (2) In this journal in 1985, Heath considered a section of an idealized snail shell; altering the overlap of adjacent whorls alters the ratio of shell material to volume enclosed. Heath located an overlap that minimized perimeter/area of the cross-section, which is different from minimizing surface-area/volume. I derive the surface area of a logarithmic helicospiral; some formulae used previously are slightly incorrect. Heath's alteration of overlap changed shell volume and aperture area; a more meaningful reanalysis keeps these characters constant. (3) In 1987, Fitter used a two-dimensional model of plant roots to show that topology, growth rate, and nutrient diffusion rate affect exploitation efficiency (area of nutrient depletion zone/volume of root). Recalculation and reanalysis show that only part of the effect of topology depends on overlap of depletion zones. I compare Fitter's model to coplanar root systems with three-dimensional depletion zones and then to fully three-dimensional branching. Finally I discuss further examples in which two-dimensional models could mislead.     

Robustness of size–structure across ecological networks in pelagic systems

The study of biomass size distributions has become an important tool for addressing aquatic ecosystem complexity and the consequences of anthropogenic disturbances. However, it remains unclear how changes in pelagic food web topology affect the biomass size–structure. Employing a dynamic multispecies bioenergetic consumer-resource model, we simulated biomass trajectories over time in 10,000 virtual networks of varying topology to address which food web properties are important in determining size–structure in pelagic systems. The slopes of the normalized biomass size spectra (NBSS) and Pareto’s shape parameter (γ) of our modeled communities are consistent with theoretically expected values for steady-state systems and empirical values reported for several aquatic ecosystems. We found that the main drivers of the NBSS slope and Pareto’s γ were the slope of the relationship between body mass and trophic level, the maximum trophic level of the food web, and the stability of total community biomass. Our analyses showed a clear conservative trend in pelagic community size–structure as demonstrated by the robustness of the NBSS slope and Pareto’s γ against most of the topological changes in virtual networks. Nevertheless, these analyses also caution that major disturbances in large-bodied or top-trophic level individuals may disrupt this stable pattern.     

WBE 模型及其在生态学中的应用：研究概述

介绍了WBE模型,综述了该模型在生态学中的应用进展。WBE模型,以及以该模型为基础的MTE模型,假设生物体为自相似分形网络结构,提出代谢速率和个体大小之间存在3/4指数关系,分别预测了从个体到生物圈多个尺度上的生物属性之间的异速生长关系,而且部分得到了验证。WBE模型的应用涵盖了个体组织生物量、年生长率,种群密度和生态系统单位面积产量、能量流动率等多个方面;即使在生物圈大尺度上,WBE模型也可用来预测试验中无法直接测量的特征变量的属性,如全球碳储量的估算等。至今,关于WBE和MTE模型仍然存在各种褒贬争论,讨论焦点主要集中于模型建立的前提假设以及权度指数的预测。今后的研究工作应规范试验技术和方法,考虑物种多样性和环境等因素的影响,提出符合各类生物的模型结构体系,使其具有更广泛的应用性和预测性。     

Kinetic modeling of cellulosic biomass to ethanol via simultaneous saccharification and fermentation: Part II. Experimental validation using waste paper sludge and anticipation of CFD analysis

A kinetic model of cellulosic biomass conversion to ethanol via simultaneous saccharification and fermentation (SSF) developed previously was validated experimentally using paper sludge as the substrate. Adsorption parameters were evaluated based on the data obtained at various values for fractional cellulose conversion. The adsorption model was then combined with batch SSF data to evaluate the cellulose hydrolysis parameters. With the parameters evaluated for the specific substrate, the discrete model was able to predict SSF successfully both with discrete addition of cellulase only and with discrete feeding of substrate, cellulase, and media. The model tested in this study extends the capability of previous SSF models to semi-continuous feeding configurations, and invites a mechanistic interpretation of the recently observed trend of increasing conversion with decreasing feeding frequency [Fan et al. (2007a) Bioprocess Biosyst Eng 30(1):27-34]. Our results also support the feasibility and utility of determining adsorption parameters based on data obtained at several conversions, particularly when the model is to be applied to extended reaction times rather than only initial hydrolysis rates. The revised model is considerably more computationally efficient than earlier models, and appears for many conditions to be within the capability of simulation using computational fluid dynamics.     

Food webs: Ordering species according to body size yields high degree of intervality

Food webs, the networks describing “who eats whom” in an ecosystem, are nearly interval, i.e. there is a way to order the species so that almost all the resources of each consumer are adjacent in the ordering. This feature has important consequences, as it means that the structure of food webs can be described using a single (or few) species' traits. Moreover, exploiting the quasi-intervality found in empirical webs can help build better models for food web structure. Here we investigate which species trait is a good proxy for ordering the species to produce quasi-interval orderings. We find that body size produces a significant degree of intervality in almost all food webs analyzed, although it does not match the maximum intervality for the networks. There is also a great variability between webs. Other orderings based on trophic levels produce a lower level of intervality. Finally, we extend the concept of intervality from predator-centered (in which resources are in intervals) to prey-centered (in which consumers are in intervals). In this case as well we find that body size yields a significant, but not maximal, level of intervality. These results show that body size is an important, although not perfect, trait that shapes species interactions in food webs. This has important implications for the formulation of simple models used to construct realistic representations of food webs.     

Trait-dependency of trophic interactions in zooplankton food webs

Anthropogenic change in the abundance or identity of dominant top predators may induce reorganizations in whole food webs. Predicting these reorganizations requires identifying the biological rules that govern trophic niches. However, we still lack a detailed understanding of the respective contributions of body size, behaviour (e.g. match between predator hunting mode and prey antipredator strategy), phylogeny and/or ontogeny in determining both the presence and strength of trophic interactions. Here, we address this question by measuring zooplankton numerical response to fish predators in lake enclosures. We compared the fit to zooplankton count data of models grouping zooplankters based either on 1) body sizes, 2) antipredator behaviour, 3) body size combined with antipredator behaviour or on 4) phylogeny combined with ontogeny (i.e. different life stages of copepods). Body size was a better predictor of zooplankton numerical response to fish than antipredator behaviour, but combining body size and behaviour provided even better predictions. Models based on phylogeny combined with ontogeny clearly outperformed those based on other zooplankton grouping rules, except when phylogeny was poorly resolved. Removing ontogenetic information plagued the predictive power of the highly-resolved (genus-level) phylogenetic grouping but not of medium-resolved or poorly-resolved phylogenetic grouping. Our results support the recent use of phylogeny as a superior surrogate for traits controlling trophic niches, and further highlight the added value of combining phylogeny with ontogenetic traits. Further improvements in our mechanistic understanding of how trophic networks are shaped are bound to uncovering the trophic traits captured by phylogeny and ontogeny, but that currently remain hidden to us.     

Prey size diversity hinders biomass trophic transfer and predator size diversity promotes it in planktonic communities

Body size exerts multiple effects on plankton food-web interactions. However, the influence of size structure on trophic transfer remains poorly quantified in the field. Here, we examine how the size diversity of prey (nano-microplankton) and predators (mesozooplankton) influence trophic transfer efficiency (using biomass ratio as a proxy) in natural marine ecosystems. Our results support previous studies on single trophic levels: transfer efficiency decreases with increasing prey size diversity and is enhanced with greater predator size diversity. We further show that communities with low nano-microplankton size diversity and high mesozooplankton size diversity tend to occur in warmer environments with low nutrient concentrations, thus promoting trophic transfer to higher trophic levels in those conditions. Moreover, we reveal an interactive effect of predator and prey size diversities: the positive effect of predator size diversity becomes influential when prey size diversity is high. Mechanistically, the negative effect of prey size diversity on trophic transfer may be explained by unicellular size-based metabolic constraints as well as trade-offs between growth and predation avoidance with size, whereas increasing predator size diversity may enhance diet niche partitioning and thus promote trophic transfer. These findings provide insights into size-based theories of ecosystem functioning, with implications for ecosystem predictive models.     

Division synchrony and the dynamics of microbial populations: A size-specific model

A discrete, environmentally coupled, size-specific model of microbial population dynamics in continuous culture is presented. It is mathematically simpler than other models based on similar assumptions and lends itself to numerical and analytic solutions. It displays several phenomena which have been reported in the experimental literature but which are not well understood; specifically, a loose relationship between biomass and numbers (i.e., a time lag between mass growth and cell division) and a critical damping of biomass while numbers continue to oscillate. In addition, the model provides several new predictions: The stable biomass distribution is independent of the environmental factors considered in the model and uniformly distributes the biomass among the size classes. The rate of approach to stability and the frequency of waves through the size distributions are a function of the flow rate and the variance in rate of growth and size at division. The model should provide a useful basis for studying the effects of size specificity on the dynamics of microbial populations cultured in chemostats.     

Modeling the growth of individuals in plant populations: local density variation in a strand population of Xanthium strumarium (Asteraceae)

We studied the growth of individual Xanthium strumarium plants growing at four naturally occurring local densities on a beach in Maine: (1) isolated plants, (2) pairs of plants ≤1 cm apart, (3) four plants within 4 cm of each other, and (4) discrete dense clumps of 10-39 plants. A combination of nondestructive measurements every 2 wk and parallel calibration harvests provided very good estimates of the growth in aboveground biomass of over 400 individual plants over 8 wk and afforded the opportunity to fit explicit growth models to 293 of them. There was large individual variation in growth and resultant size within the population and within all densities. Local crowding played a role in determining plant size within the population: there were significant differences in final size between all densities except pairs and quadruples, which were almost identical. Overall, plants growing at higher densities were more variable in growth and final size than plants growing at lower densities, but this was due to increased variation among groups (greater variation in local density and/or greater environmental heterogeneity), not to increased variation within groups. Thus, there was no evidence of size asymmetric competition in this population. The growth of most plants was close to exponential over the study period, but half the plants were slightly better fit by a sigmoidal (logistic) model. The proportion of plants better fit by the logistic model increased with density and with initial plant size. The use of explicit growth models over several growth intervals to describe stand development can provide more biological content and more statistical power than "growth-size" methods that analyze growth intervals separately.     

A Generic View of Classic Microbial Growth Models

General theoretical aspects are reviewed of models for microbial growth and endogenous metabolism. The focus is on a generic cell model with two components. Growth is represented as the increase of one of these components (the structural scaffolding or 'frame'). A novel feature of the present generic model is the explicit modelling of (partial) metabolic shutdown under conditions where maintenance requirements cannot be met.Two different approaches to mechanistic underpinnings for the classic models are outlined. The first approach is based on a bimolecular reaction between the non-permanent biomass component and the permanent biomass component. The second approach is based on cellular control systems.     

Body size and food web structure: testing the equiprobability assumption of the cascade model

The cascade model successfuly predicts many patterns in reported food webs. A key assumption of this model is the existence of a predetermined trophic hierarchy; prey are always lower in the hierarchy than their predators. At least three studies have suggested that, in animal food webs, this hierarchy can be explained to a large extent by body size relationships. A second assumption of the standard cascade model is that trophic links not prohibited by the hierarchy occur with equal probability. Using nonparametric contingency table analyses, we tested this ”equiprobability hypothesis” in 16 published animal food webs for which the adult body masses of the species had been estimated. We found that when the hierarchy was based on body size, the equiprobability hypothesis was rejected in favor of an alternative, ”predator-dominance” hypothesis wherein the probability of a trophic link varies with the identity of the predator. Another alternative to equiprobabilty is that the probability of a trophic link depends upon the ratio of the body sizes of the two species. Using nonparametric regression and liklihood ratio tests, we show that a size-ratio based model represents a significant improvement over the cascade model. These results suggest that models with heterogeneous predation probabilities will fit food web data better than the homogeneous cascade model. They also suggest a new way to bridge the gap between static and dynamic food web models. Received: 3 February 1999 / Accepted: 26 October 1999     

春季长江口邻近外海网采浮游生物的生物量谱

对2005年春季黄海南部、东海北部近江口外海水域网采的浮游生物个体大小的粒径分布进行研究,确定各粒级大小的功能群组成,建立此季该调查水域网采浮游生物的生物量谱.样品是用小、中、大型浮游生物网(网孔径为77、160、 505μm)采集所得.三网具采集浮游生物个体大小粒径是连续的,其中浮游植物,其等效粒径(ESD)和含碳量范围分别为5～250μm、15pg～146ng;浮游动物,含碳量范围为115ng～7.5mg,ESD分别为120μm～5.8mm、200μm～2cm.所得网采浮游生物的标准生物量谱,总测区的斜率为-0.607±0.059、截距为19.45±0.46;各站位的生物量谱斜率为-0.889～-0.455、截距为12.866～16.863,两特征参数分布规律为南高北低,且具有显著的站位间差异.相关性分析表明截距和回归系数与粒级多样性有关.     

Testing mechanistic models of growth in insects

Insects are typified by their small size, large numbers, impressive reproductive output and rapid growth. However, insect growth is not simply rapid; rather, insects follow a qualitatively distinct trajectory to many other animals. Here we present a mechanistic growth model for insects and show that increasing specific assimilation during the growth phase can explain the near-exponential growth trajectory of insects. The presented model is tested against growth data on 50 insects, and compared against other mechanistic growth models. Unlike the other mechanistic models, our growth model predicts energy reserves per biomass to increase with age, which implies a higher production efficiency and energy density of biomass in later instars. These predictions are tested against data compiled from the literature whereby it is confirmed that insects increase their production efficiency (by 24 percentage points) and energy density (by 4 J mg⁻¹) between hatching and the attainment of full size. The model suggests that insects achieve greater production efficiencies and enhanced growth rates by increasing specific assimilation and increasing energy reserves per biomass, which are less costly to maintain than structural biomass. Our findings illustrate how the explanatory and predictive power of mechanistic growth models comes from their grounding in underlying biological processes.     

Using successional theory to measure marine ecosystem health

Marine ecosystems are diverse and complex, providing significant challenges to the development of generalizable metrics of ecosystem health. Of particular concern is the varied form of change caused by multiple human activities, which limits the capacity to generate a single measure to encapsulate the overall condition of the ecosystem. Here we consider how successional theory can help to simplify our understanding of marine community structure, especially when viewed in context of human disturbance. During succession, the emergent properties of communities change in predictable ways. As communities mature, there is an increase in total production and biomass, the mean size of organisms, the level of internal recycling of food and nutrients, and the mean trophic level. Using a set of multi-species trophic models, we explore the changes in community structure that are likely to occur during succession. These changes include increases in biomass within trophic levels due to decreased rates of energy and food loss through trophic and production inefficiencies, and potential shifts from top-down control early in succession to bottom-up later. Because human activities disproportionately favor early-successional species, we can gain insights by considering community degradation in the context of succession being played in reverse. Indicators of health based on ecological succession thus provide a mechanistic view to measure the impact of human activities (both positive and negative) on marine ecosystems.     

Does the energy equivalence rule apply to intertidal macrobenthic communities?

Energy equivalence assumes equal contribution of large and small species to production and energy flow in communities. As in a double logarithmic plot, physiological rates decline with body weight by –0.25, log biomass should increase by 0.25 and log abundance decline by –0.75 with log species weight, when this concept is valid. This was tested with annual data sets of the macrobenthos of 4 intertidal sites in the German Wadden Sea (Königshafen) and 3 sites in a south Portuguese lagoon (Ria Formosa). Only abundance data from two of these sites displayed significantly negative slopes with mean body size of the species. Biomass and secondary production data were significantly positively correlated with mean body size for all Ria Formosa sites and also for the biomass of a mussel bed in Königshafen. However, high variation in body size of the individuals of a species limits interpretation of these plots.It is preferable to test this concept by body weight classes regardless of its species composition. At Königshafen, biomass and production displayed two distinct peaks. One peak at small body size was caused by browsing species. The other peak at larger body size was caused by animals which potentially extract their food from the water column. This bimodality was only vaguely reflected at one station in the Ria Formosa, possibly because of a dominance of detritus feeding species. In a normalized form (log biomass or production / width of size classvs. log size class), these spectra imply a dominance of small individuals in biomass and production at all sites (except for a mussel bank at Königshafen). This is interpreted as a consequence of permanent disturbances.     

Some interrelationships among the regression coefficient estimates arising in a class of models appropriate to response-time data.

Interrelationships among three response-time models which incorporate covariate information are explored. The most general of these models is the logistic-exponential in which the log odds of the probability of responding in a fixed interval is assumed to be a linear function of the covariates; this model includes a parameter W for the width of discrete time intervals in which responses occur. As W leads to O this model is equivalent to a continuous time exponential model in which the log hazard is linear in the covariates. As W leads to infininity it is equivalent to a continuous time exponential model in which the hazard itself is a linear function of the covariates. This second model was fitted to the data used in an earlier publication describing the logistic exponential model, and very close agreement of the estimates of the regression coefficients is demonstrated.