Successional theories

Succession is a fundamental concept in ecology because it indicates how species populations, communities, and ecosystems change over time on new substrate or after a disturbance. A mechanistic understanding of succession is needed to predict how ecosystems will respond to land-use change and to design effective ecosystem restoration strategies. Yet, despite a century of conceptual advances a comprehensive successional theory is lacking. Here we provide an overview of 19 successional theories (‘models’) and their key points, group them based on conceptual similarity, explain conceptual development in successional ideas and provide suggestions how to move forward. Four groups of models can be recognised. The first group (patch & plants) focuses on plants at the patch level and consists of three subgroups that originated in the early 20th century. One subgroup focuses on the processes (dispersal, establishment, and performance) that operate sequentially during succession. Another subgroup emphasises individualistic species responses during succession, and how this is driven by species traits. A last subgroup focuses on how vegetation structure and underlying demographic processes change during succession. A second group of models (ecosystems) provides a more holistic view of succession by considering the ecosystem, its biota, interactions, diversity, and ecosystem structure and processes. The third group (landscape) considers a larger spatial scale and includes the effect of the surrounding landscape matrix on succession as the distance to neighbouring vegetation patches determines the potential for seed dispersal, and the quality of the neighbouring patches determines the abundance and composition of seed sources and biotic dispersal vectors. A fourth group (socio-ecological systems) includes the human component by focusing on socio-ecological systems where management practices have long-lasting legacies on successional pathways and where regrowing vegetations deliver a range of ecosystem services to local and global stakeholders. The four groups of models differ in spatial scale (patch, landscape) or organisational level (plant species, ecosystem, socio-ecological system), increase in scale and scope, and reflect the increasingly broader perspective on succession over time. They coincide approximately with four periods that reflect the prevailing view of succession of that time, although all views still coexist. The four successional views are: succession of plants (from 1910 onwards) where succession was seen through the lens of species replacement; succession of communities and ecosystems (from 1965 onwards) when there was a more holistic view of succession; succession in landscapes (from 2000 onwards) when it was realised that the structure and composition of landscapes strongly impact successional pathways, and increased remote-sensing technology allowed for a better quantification of the landscape context; and succession with people (from 2015 onwards) when it was realised that people and societal drivers have strong effects on successional pathways, that ecosystem processes and services are important for human well-being, and that restoration is most successful when it is done by and for local people. Our review suggests that the hierarchical successional framework of Pickett is the best starting point to move forward as this framework already includes several factors, and because it is flexible, enabling application to different systems. The framework focuses mainly on species replacement and could be improved by focusing on succession occurring at different hierarchical scales (population, community, ecosystem, socio-ecological system), and by integrating it with more recent developments and other successional models: by considering different spatial scales (landscape, region), temporal scales (ecosystem processes occurring over centuries, and evolution), and by taking the effects of the surrounding landscape (landscape integrity and composition, the disperser community) and societal factors (previous and current land-use intensity) into account. Such a new, comprehensive framework could be tested using a combination of empirical research, experiments, process-based modelling and novel tools. Applying the framework to seres across broadscale environmental and disturbance gradients allows a better insight into what successional processes matter and under what conditions.     

A hierarchical consideration of causes and mechanisms of succession

Questions of successional pattern and causality have been central concerns in vegetation ecology. In this paper we address the limits of the overextended models of Connell and Slatyer by discussing problems encountered in field tests. To help prevent such problems, we define the essential concepts needed to understand succession: pathway, cause, mechanism, and model. We then suggest a more complete enumeration of successional causes, and place them in a three-level hierarchy. The highest level in the hierarchy defines the general and universal conditions under which succession occurs: (1) availability of open sites, (2) differential availability of species, and (3) differential performance of species at the site. To provide a more detailed under-standing of succession, each of these causes is decomposed into ecological processes. A further decomposition results in the third level of the hierarchy, which is required to elucidate the mechanisms of succession at particular sites and to make detailed predictions. The hierarchy allows the appropriate causes to be chosen to answer questions about succession at the desired level of generality or level of organization. Recognizing the appropriate level(s) in the hierarchy is critical for the successful explanation of succession, design of experiments, statement of predictions, construction of models and development of general theory.We thank Prof. Joseph Connell for comments and helpful discussion and Prof. F. A. Bazzaz, Dr. P. S. White, Dr. L. R. Walker and the members of the Plant Strategy and Vegetation Dynamics Lab. at Rutgers for criticism. Preparation of this paper was supported by the Mary Flagler Charitable Trust through the Institute of Ecosystem Studies.     

陆地植物群落物种多样性的梯度变化特征

研究陆地植物落物种多样性随环境因子及群落演替梯度的变化特征是揭示生物多样性与生态因子相互关系的重要方面,根据近期国内外的文献,综述了这方面的研究进展。随纬度的降低,通常物咱多样性随中,随不分梯度的变化,物种多样性的变化有６种趋势;随海拔高的变化,物处多样性有５种模式;随土壤养分梯度的变化,表现出不同的规律;演替过程中物种多样生的变化趋势相似。关于植物群落物种多样性梯度格局的机制有多种假说,但仍需进     

Quantifying ecological memory during forest succession: A case study from lower subtropical forest ecosystems in South China

The concept of ecological memory provides a new perspective for research on forest succession by including historical factors and the initial state of ecological processes. However, there are still significant gaps between the concept and its application. We selected nine proxy indicators (plant species, soil seed banks, soil microbes, soil animals, birds, soil age, soil pollen, soil mineral distribution, and light environment) and developed a method to quantify ecological memory and succession in a subtropical forest succession in South China. Taking the climax-monsoon evergreen broad-leaved forest as the reference ecosystem, we found that ecological memory increased nonlinearly and accumulated following a specific assembly rule during succession. Memory concerning major soil microbes and soil animals, which improve the soil substrate, mainly accumulated from the initial to the early successional stage. Memory concerning the number of bird species and the availability of light, which ensure a source of regenerative seeds and the survival of understory seedlings, mainly accumulated from the early to middle successional stages. Memory concerning vegetation and soil seed banks mainly accumulated late in succession, guaranteeing that the ecosystem would reach the regional climax stage. Prospective memory was greater than retrospective memory in every successional stage except the late stage, which indicated that all stages but the late stage were undergoing progressive succession. Our study demonstrates that the concept of ecological memory and the proposed evaluation framework are useful for guiding research on succession and restoration, and especially for assessing how “far” a restored ecosystem is from a reference ecosystem or how far a restored ecosystem has deviated from its natural succession trajectory.     

Mort ou persistance du darwinisme ? Regard d’un épistémologue

The article examines why evolutionary biologists have been haunted by the question whether they are “Darwinian” or “non-Darwinian” ever since Darwin's Origin of species. Modern criticisms addressed to Darwinism are classified into two categories: those concerning Darwin's hypothesis of “descent with modification” and those addressed to the hypothesis of natural selection. In both cases, although the particular models that Darwin proposed for these two hypotheses have been significantly revised and expanded, Darwin's general framework has constrained and canalized evolutionary research, in the sense that it has settled an array of possible theoretical choices. Gould's changing attitudes regarding Darwinism is taken as a striking illustration of this interpretation.     

Landscape Pattern Dynamics and Mechanisms during Vegetation Restoration: A Multiscale,Hierarchical Patch Dynamics Approach

The spatial pattern of vegetation changes during ecological restoration, and these changes are affected by the process of restoration. The objective of this study was to integrate the pattern and mechanism of forest restoration in the Dinghushan Nature Reserve (DNR), Guangdong, China, based on data from remote sensing and long‐term field observations. We studied the pattern dynamics of three main forest types and their underlying mechanisms during restoration following a multiscale, hierarchical patch dynamics framework that integrates population, community, and landscape processes. Remote sensing data were used to determine the changes in landscape pattern during different periods of forest restoration from 1978 to 2006. At the landscape scale, the number, area, and perimeter of the needle/broad‐leaved mixed forest (MF) and the evergreen broad‐leaved forest (BF) increased, whereas those of the tropical needle‐leaved forest (NF) decreased during succession. Our analysis based on long‐term field observations indicated that the change rate of NF was lower than that of MF during 1981–1996, but became much higher during 1996–2007. The rate of change in landscape pattern and the progression of succession stages were consistent with each other. Our results also showed that species regeneration and community succession are the biological basis of forest landscape dynamics during vegetation restoration. Landscape pattern analysis allowed us to show “what” happened during vegetation restoration and “where,” and population and community analysis indicated “why” and “how” it happened.     

Marine invertebrates in an algal succession. II. Tests of hypotheses to explain changes in diversity with succession

The invertebrate community living in algal mats on intertidal boulders was studied for 2 yr. The diversity and abundance of the animals increased between the early and middle stages of algal succession, then remained similar into the later stage. Three possible mechanisms producing this pattern were investigated experimentally by manipulating natural algal mats and plastic algal mimics in the field and laboratory and evaluating the community of colonizing invertebrates. The first, the “ecological time” hypothesis, suggests that there are more species and individuals in later stages because they accumulate slowly with time; this hypothesis was tested experimentally and rejected. A second, “algal toxicity” hypothesis suggests that species richness and abundance are lower in earlier successional stages because the early colonizing green algae are more toxic to animals than are the later red algae. This hypothesis was also tested experimentally and rejected. The third, “habitat complexity” hypothesis suggests that increases in complexity of physical aspects of algal structure (biomass, surface area) cause increases in invertebrate richness and abundance. The fact that this result was found in both living algae and plastic mimics indicates that biological aspects of algal structure apparently have only minor importance. Algal biomass and surface area increase from early to middle successional stages; middle and late successional stages are similar. In general, increases in these physical aspects of algal structure produce concomitant increases in the abundance and diversity of the associated animal community. With higher biomass and surface area, increased numbers of individuals accumulate in algal mats. Because a larger sample of the available pool of individuals is therefore collected, more species are found in a given area of algal mat when the structure is more complex. The successional patterns of increase in species richness of this invertebrate community seem to result from this sampling phenomenon, rather than from increases in numbers of resources (i.e., “niches”).     

The evolution and ecology of multiple antipredator defences

Prey seldom rely on a single type of antipredator defence, often using multiple defences to avoid predation. In many cases, selection in different contexts may favour the evolution of multiple defences in a prey. However, a prey may use multiple defences to protect itself during a single predator encounter. Such “defence portfolios” that defend prey against a single instance of predation are distributed across and within successive stages of the predation sequence (encounter, detection, identification, approach (attack), subjugation and consumption). We contend that at present, our understanding of defence portfolio evolution is incomplete, and seen from the fragmentary perspective of specific sensory systems (e.g., visual) or specific types of defences (especially aposematism). In this review, we aim to build a comprehensive framework for conceptualizing the evolution of multiple prey defences, beginning with hypotheses for the evolution of multiple defences in general, and defence portfolios in particular. We then examine idealized models of resource trade-offs and functional interactions between traits, along with evidence supporting them. We find that defence portfolios are constrained by resource allocation to other aspects of life history, as well as functional incompatibilities between different defences. We also find that selection is likely to favour combinations of defences that have synergistic effects on predator behaviour and prey survival. Next, we examine specific aspects of prey ecology, genetics and development, and predator cognition that modify the predictions of current hypotheses or introduce competing hypotheses. We outline schema for gathering data on the distribution of prey defences across species and geography, determining how multiple defences are produced, and testing the proximate mechanisms by which multiple prey defences impact predator behaviour. Adopting these approaches will strengthen our understanding of multiple defensive strategies.     

Nestedness and successional trajectories of macroinvertebrate assemblages in man-made wetlands

Current successional models, primarily those based on floral succession, propose several distinct trajectories based on the integration of two key hypotheses from succession theory: convergence versus divergence in species composition among successional sites, and progression towards versus deviation from a desired reference state. We applied this framework to faunal succession, including differential colonization between active and passive dispersers, and the nested patterns generated as a consequence of this peculiarity. Nine man-made wetlands located in three different areas, from 0–3 years from wetland creation, were assessed. In addition, 91 wetlands distributed throughout the region were used as references for natural macroinvertebrate communities. We predicted the following: (1) highly nested structures in pioneering assemblages will decrease to lower mid-term values due to a shift from active pioneering taxa to passive disperser ones; (2) passive idiosyncratic taxa will elicit divergent successional trajectories among areas; (3) the divergent trajectories will provoke lower local and higher regional diversity values in the mid-term assemblages than in pioneer assemblages. Our results were largely congruent with hypotheses (1) and (2), diverging from the anticipated patterns only in the case of the temporary wetlands area. However, overall diversity trends based on hypothesis (3) did not follow the expected pattern. The divergent successional trajectories did not compensate for regional biodiversity losses that occurred as a consequence of pioneering colonizer decline over time. Consequently, we suggest reconsidering wetland construction for mitigation purposes within mid-term time frames (≤3 years). Wetlands may not offset, within this temporal scenario, regional biodiversity loss because the ecosystem may not support idiosyncratic taxa from natural wetlands.     

A Historical Perspective for the Catalytic Reaction Mechanism of Glycosidase; So As to Bring about Breakthrough in Confusing Situation

Various catalytic reaction models have been proposed as the reaction mechanisms of glycosidases, but a reasonable and unitary model capable of interpreting both “inverting” and “retaining” glycosidase reactions remains to be established. As for the models proposed to date, the nucleophilic displacement mechanism and the oxocarbenium ion intermediate mechanism are widely known, but recently the former is widely accepted, and so the general tendency of world opinion appears to favor it. This reaction model, however, is considered to comprise some inconsistencies that cannot be neglected from the viewpoint of reactivity in organic chemistry. While the nucleophilic displacement mechanism is often applied to reactions of glycosidases, it appears unlikely that such reactions actually occur. This review argues that the oxocarbenium ion intermediate reaction mechanism is more rational than the nucleophilic displacement reaction mechanism, as the action mode of glycosidases and related enzymes.     

Mechanistic systems models of cell signaling networks: a case study of myocyte adrenergic regulation

Signal transduction networks coordinate a wide variety of cellular functions, including gene expression, metabolism, and cell fate processes. Understanding biological networks quantitatively is a major challenge to post-genomic biology, and mechanistic systems models will be crucial for this task. Here, we review approaches towards developing mechanistic systems models of established cell signaling networks. The ability of mechanistic system models to generate testable biological hypotheses and experimental strategies is discussed. As a case study of model development and analysis, we examined the functional roles of phospholamban, the L-type calcium channel, the ryanodine receptor, and troponin I phosphorylation upon β-adrenergic stimulation in the rat ventricular myocyte. Model analysis revealed that while protein kinase A-mediated phosphorylation of the ryanodine receptor greatly increases its calcium sensitivity, calcium autoregulation may adapt quickly by negating potential increases in contractility. Systematic combinations of in silico perturbations supported the conclusion that phospholamban phosphoregulation is the primary mechanism for increased sarcoplasmic reticulum load and calcium relaxation rate during β-adrenergic stimulation, while both phospholamban and the L-type calcium channel contribute to increased systolic calcium. Combined with detailed experimental studies, mechanistic systems models will be valuable for developing a quantitative understanding of cell signaling networks.     

Ashmole's hypothesis and the latitudinal gradient in clutch size

One enduring priority for ecologists has been to understand the cause(s) of variation in reproductive effort among species and localities. Avian clutch size generally increases with increasing latitude, both within and across species, but the mechanism(s) driving that pattern continue to generate hypotheses and debate. In 1961, a Ph.D. student at Oxford University, N. Philip Ashmole, proposed the influential hypothesis that clutch size varies in direct proportion to the seasonality of resources available to a population. Ashmole's hypothesis has been widely cited and discussed in the ecological literature. However, misinterpretation and confusion has been common regarding the mechanism that underlies Ashmole's hypothesis and the testable predictions it generates. We review the development of well-known hypotheses to explain clutch size variation with an emphasis on Ashmole's hypothesis. We then discuss and clarify sources of confusion about Ashmole's hypothesis in the literature, summarise existing evidence in support and refutation of the hypothesis, and suggest some under-utilised and novel approaches to test Ashmole's hypothesis and gain an improved understanding of the mechanisms responsible for variation in avian clutch size and fecundity, and life-history evolution in general.     

Towards a molecular understanding of cytokinesis

In this review, we focus on recent discoveries regarding the molecular basis of cleavage furrow positioning and contractile ring assembly and contraction during cytokinesis. However, some of these mechanisms might have different degrees of importance in different organisms. This synthesis attempts to uncover common themes and to reveal potential relationships that might contribute to the biochemical and mechanical aspects of cytokinesis. Because the information about cytokinesis is still fairly rudimentary, our goal is not to present a definitive model but to present testable hypotheses that might lead to a better mechanistic understanding of the process.     

Generative models versus underlying symmetries to explain biological pattern

Mathematical models play an increasingly important role in the interpretation of biological experiments. Studies often present a model that generates the observations, connecting hypothesized process to an observed pattern. Such generative models confirm the plausibility of an explanation and make testable hypotheses for further experiments. However, studies rarely consider the broad family of alternative models that match the same observed pattern. The symmetries that define the broad class of matching models are in fact the only aspects of information truly revealed by observed pattern. Commonly observed patterns derive from simple underlying symmetries. This article illustrates the problem by showing the symmetry associated with the observed rate of increase in fitness in a constant environment. That underlying symmetry reveals how each particular generative model defines a single example within the broad class of matching models. Further progress on the relation between pattern and process requires deeper consideration of the underlying symmetries.     

Mathematical modelling of ecological succesion—a review

The present review gives an account of the applicability of mathematical modelling in ecological succession studies. The ability of particular model types to solve problems of both theory and management is discussed. The Markovian models are found to be useful for short term predictions, but of very limited value for theoretical considerations. Finally, the predictability of successional pathways is discussed. It is argued that the less we understand about processes in vegetation dynamics, the more we will see the course of succession as random and unpredictable.     

Linking environmental variability and fish performance: integration through the concept of scope for activity

Investigating the biological mechanisms linking environmental variability to fish production systems requires the disentangling of the interactions between habitat, environmental adaptation and fitness. Since the number of environmental variables and regulatory processes is large, straightening out the environmental influences on fish performance is intractable unless the mechanistic analysis of the 'fish-milieu' system is preceded by an understanding of the properties of that system. While revisiting the key points in our currently poorly integrated understanding of fish ecophysiology, we have highlighted the explanatory potential contained within Fry's (Fry 1947 Univ. Toronto Stud. Biol. Ser. 55, 1-62) concept of metabolic scope and categorization of environmental factors. These two notions constitute a pair of powerful tools for conducting an external (at the emerging property level) analysis of the environmental influences on fish, as well as an internal (mechanistic) examination of the behavioural, morphological and physiological processes involved. Using examples from our own and others work, we have tried to demonstrate that Fry's framework represents a valuable conceptual basis leading to a broad range of testable ecophysiological hypotheses.     

Key evolutionary innovations and their ecological mechanisms

Explanations for taxonomic diversity in a particular clade often implicate evolutionary innovations, possessed by members of the clade, that are thought to have favoured diversification. We review such “key innovation”; hypotheses, the ecological mechanisms involved, and potential tests of such hypotheses.

Key innovation hypotheses can be supported by evidence of ecological mechanism and by comparative tests. We argue that both are necessary for convincing support. In fact, few key innovation hypotheses are currently backed by either one.

We group ecological mechanisms of diversification in three major classes. Diversification may be spurred by innovations that: I) allow invasion of new adaptive zones; II) increase fitness, allowing one clade to replace another; or III) increase the propensity for reproductive or ecological specialization. Key innovations in different classes are likely to produce different evolutionary patterns, and therefore may be supported by different kinds of ecological evidence.     

Reverse engineering the lordosis behavior circuit

Reverse engineering takes the facts we know about a device or a process and reasons backwards to infer the principles underlying the structure-function relations. The goal of this review is to apply this approach to a well-studied hormone-controlled behavior, namely the reproductive stance of female rodents, lordosis. We first provide a brief overview on the considerable amount of progress in the analysis of female reproductive behavior. Then, we propose an analysis of the mechanisms of this behavior from a reverse-engineering perspective with the goal of generating novel hypotheses about the properties of the circuitry elements. In particular, the previously proposed neuronal circuit modules, feedback signals, and genomic mechanisms are considered to make predictions in this manner. The lordosis behavior itself appears to proceed ballistically once initiated, but negative and positive hormonal feedback relations are evident in its endocrine controls. Both rapid membrane-initiated and slow genomic hormone effects contribute to the behavior's control. We propose that the value of the reverse-engineering approach is based on its ability to provide testable, mechanistic hypotheses that do not emerge from either traditional evolutionary or simple reductionistic perspectives, and several are proposed in this review. These novel hypotheses may generalize to brain functions beyond female reproductive behavior. In this way, the reverse-engineering perspective can further develop our conceptual frameworks for behavioral and systems neuroscience.