Progress in wetland restoration ecology

It takes more than water to restore a wetland. Now, scientists are documenting how landscape setting, habitat type, hydrological regime, soil properties, topography, nutrient supplies, disturbance regimes, invasive species, seed banks and declining biodiversity can constrain the restoration process. Although many outcomes can be explained post hoc, we have little ability to predict the path that sites will follow when restored in alternative ways, and no insurance that specific targets will be met. To become predictive, bolder approaches are now being developed, which rely more on field experimentation at multiple spatial and temporal scales, and in many restoration contexts.     

Alternative logical states

N. W. H. Mason, J. B. Wilson and J. B. Steel argue that there is no logical conceptual basis, and no empirical data, to support an association between environmental adversity and the occurrence of alternative stable states. While we agree that robust debate on the relative frequency of occurrence of alternative stable states is valuable, any apparent logic in their argument is confounded by misinterpretation and direct misrepresentation of our earlier arguments ( Didham et al. 2005 ), despite the fact that we clarified many of the same issues in response to T. Fukami and W. G. Lee ( Didham and Norton 2006 ). Opinion, not logic or evidence, underlies Mason, Wilson and Steel's argument, and this does little to further our understanding of why some systems exhibit alternative stable states that are resilient to restoration management efforts.     

Belowground positive and negative feedbacks on CO2 growth enhancement

In this paper we present a conceptual model of integrated plant-soil interactions which illustrates the importance of identifying the primary belowground feedbacks, both positive and negative, which can simultaneously affect plant growth responses to elevated CO₂. The primary negative feedbacks share the common feature of reducing the amount of nutrients available to plants. These negative feedbacks include increased litter C/N ratios, and therefore reduced mineralization rates, increased immobilization of available nutrients by a larger soil microbial pool, and increased storage of nutrients in plant biomass and detritus due to increases in net primary productivity (NPP). Most of the primary positive feedbacks share the common feature of being plant mediated feedbacks, the only exception being Zak et al.'s hypothesis that increased microbial biomass will be accompanied by increased mineralization rates. Plant nutrient uptake may be increased through alterations in root architecture, physiology, or mycorrhizal symbioses. Further, the increased C/N ratios of plant tissue mean that a given level of NPP can be achieved with a smaller supply of nitrogen.Identification of the net plant-soil feedbacks to enhanced productivity with elevated CO₂ are a critical first step for any ecosystem. It is necessary, however, that we first identify how universally applicable the results are from one study of one ecosystem before ecosystem models incorporate this information. The effect of elevated CO₂ on plant growth (including NPP, tissue quality, root architecture, mycorrhizal symbioses) can vary greatly for different species and environmental conditions. Therefore it is reasonable to expect that different ecosystems will show different patterns of interacting positive and negative feedbacks within the plant-soil system. This inter-ecosystem variability in the potential for long-term growth responses to rising CO₂ levels implies that we need to parameterize mechanistic models of the impact of elevated CO₂ on ecosystem productivity using a detailed understanding of each ecosystem of interest.     

Alternative states in shallow lakes

Ecology of Shallow Lakes by Marten Scheffer Chapman and Hall (Population and Community Biology Series 22), 1998. ￡45.00/$79.65 hbk (xx+357 pages) ISBN 0 412 74920 3.     

Two independent positive feedbacks and bistability in the Bcl-2 apoptotic switch

Background

The complex interplay between B-cell lymphoma 2 (Bcl-2) family proteins constitutes a crucial checkpoint in apoptosis. Its detailed molecular mechanism remains controversial. Our former modeling studies have selected the ‘Direct Activation Model’ as a better explanation for experimental observations. In this paper, we continue to extend this model by adding interactions according to updating experimental findings.

Methodology/Principal Findings

Through mathematical simulation we found bistability, a kind of switch, can arise from a positive (double negative) feedback in the Bcl-2 interaction network established by anti-apoptotic group of Bcl-2 family proteins. Moreover, Bax/Bak auto-activation as an independent positive feedback can enforce the bistability, and make it more robust to parameter variations. By ensemble stochastic modeling, we also elucidated how intrinsic noise can change ultrasensitive switches into gradual responses. Our modeling result agrees well with recent experimental data where bimodal Bax activation distributions in cell population were found.

Conclusions/Significance

Along with the growing experimental evidences, our studies successfully elucidate the switch mechanism embedded in the Bcl-2 interaction network and provide insights into pharmacological manipulation of Bcl-2 apoptotic switch as further cancer therapies.     

Coupled positive feedbacks provoke slow induction plus fast switching in apoptosis

Apoptosis is a form of a programmed cell death for multicellular organisms to remove unwanted or damaged cells. This critical choice of cellular fate is an all-or-none process, but its dynamics remains unraveled. The switch-like apoptotic decision has to be reliable, and once a pro-apoptotic fate is determined it requires fast and irreversible execution. One of the key regulators in apoptosis is caspase-3. Interestingly, activated caspase-3 quickly executes apoptosis, but it takes considerable time to activate it. Here, we have analyzed this "slow induction plus fast switching" mechanism of caspase-3 through mathematical modeling and computational simulation. First, we have shown that two positive feedbacks, composed of caspase-8 and XIAP, are essential for the "slow induction plus fast switching" behavior of caspase-3. Second, we have found that XIAP in the feedback loops primarily regulates induction time of caspase-3. In many cancer cells activation of caspase-3 is suppressed. Our results suggest that reinforcement of the positive feedback by XIAP, which relieves XIAP-mediated caspase-3 inhibition, might favor a pro-apoptotic cellular fate.     

Habitat complexity and consumer-mediated positive feedbacks on a Caribbean coral reef

Theoretical and empirical evidence suggest that positive feedbacks can increase resilience in ecological communities. On Caribbean coral reefs, there have been striking shifts from physically complex communities with high coral cover to relatively homogenous communities dominated by macroalgae, which have persisted for decades. However, little is known about positive feedbacks that may maintain coral reef community states. Here, I explore a potential consumer-mediated feedback on a Jamaican reef by examining how grazing by a keystone herbivore ( Diadema antillarum ) is enhanced by physical structure, which offer refugia from predation. Surveys revealed that habitat complexity and Diadema density were positively related. Increasing habitat complexity by adding physical structure significantly decreased macroalgal cover and increased the proportion of urchins in algal habitats in field manipulations. Experimental increases in urchin density also decreased macroalgal cover, but did not affect the proportion of urchins in algal habitats. These results suggest that the low habitat complexity of macroalgal-dominated reefs may inhibit an urchin-mediated shift to coral dominance and that positive feedbacks must be considered in reef restoration efforts.     

Interlinked mutual inhibitory positive feedbacks induce robust cellular memory effects

Mutual inhibitory positive feedback (MIPF), or double-negative feedback, is a key regulatory motif of cellular memory with the capability of maintaining switched states for transient stimuli. Such MIPFs are found in various biological systems where they are interlinked in many cases despite a single MIPF can still realize such a memory effect. An intriguing question then arises about the advantage of interlinking MIPFs instead of exploiting an isolated single MIPF to realize the memory effect. We have investigated the advantages of interlinked MIPF systems through mathematical modeling and computer simulations. Our results revealed that interlinking MIPFs expands the parameter range of achieving the memory effect, or the memory region, thereby making the system more robust to parameter perturbations. Moreover, the minimal duration and amplitude of an external stimulus required for off-to-on state transition are increased and, as a result, external noises can more effectively be filtered out. Hence, interlinked MIPF systems can realize more robust cellular memories with respect to both parameter perturbations and external noises. Our study suggests that interlinked MIPF systems might be an evolutionary consequence acquired for a more reliable memory effect by enhancing robustness against noisy cellular environments.     

Islands of fertility induce co-occurring negative and positive plant-soil feedbacks promoting coexistence

Positive plant-soil feedback by “ecosystem engineers” is an important driver for the structuring and organization of resource-limited ecosystems. Although ample evidence demonstrates that plant-soil feedbacks can range from positive to strongly negative, their co-occurrence in plant communities have not yet been investigated. We test the hypothesis that the plant-soil feedback generated by the nitrogen-fixer shrub Medicago marina during primary succession in a sand dune community has a positive effect on the coexisting grass Lophochloa pubescens and a negative effect on the shrub species itself. We conducted field measurements and laboratory bioassays to evaluate (1) the effects of islands of fertility on the recruitment and growth of its ecosystem engineer and on the performance of a coexisting species and (2) the mechanisms involved that can explain the opposite effects of islands of fertility on coexisting species. Islands of fertility were present under Medicago crowns evidenced by higher available nitrogen, extractable phosphorus and potassium, organic matter, microbial activity, water holding capacity, soil humidity, and lower salt concentrations. The effects of these islands of fertility were clearly species-specific, with a facilitative impact on Lophochloa and a negative effect on Medicago recruitment. Lophochloa was denser and produced more biomass when rooted inside as compared to outside the crown area of the shrub. Contrarily, the number of seedlings of Medicago was lower inside, despite the higher seed abundance, and higher outside the crown area of adult shrubs as compared to predictions based on random distribution, thus showing a Janzen-Connell distribution. Laboratory experiments demonstrate the occurrence of Medicago negative plant-soil feedback, and that the auto-toxicity of the aboveground senescent plant material is a potentially important underlying mechanism explaining this negative feedback and the resulting Janzen-Connell distribution in the field.     

Alternative dynamical states in stage-structured consumer populations

The population dynamics of a consumer population with an internal structure is investigated. The population is divided into juvenile and adult individuals that consume different resources and do not interfere with each other. Over a broad range of external conditions (varying mortality and different resource levels), alternative stable states exist. These population states correspond to domination of juveniles and domination of adults, respectively. When mortality is varied, hysteresis between the alternative states only occurs if juveniles have more resources than adults. In the opposite case the juvenile-dominated state is stable for all values of mortality, but the adult-dominated state is not. When the population is modelled with more than one juvenile stage, the adult-dominated state becomes a periodic orbit due to a delay in the regulatory mechanism of the population dynamics. It is shown numerically that the stage-structured model converges to a model with continuous size structure for very large numbers of successive juvenile stages.     

Alternative stable states and regional community structure

Many models of local species interactions predict the occurrence of priority effects due to alternative stable equilibria (ASE). However, few empirical examples of ASE have been shown. One possible explanation for the disparity is that local ASE are difficult to maintain regionally in patch dynamic models. Here we examine two possible mechanisms for regional coexistence of species engaged in local ASE. Biotically generated heterogeneity (e.g., habitat modification that favors further invasion by conspecifics) results in regional exclusion of one species at equilibrium. In contrast, abiotic heterogeneity due to spatial variation in resource supply ratios generates local-scale ASE and ensures regional coexistence with sufficiently broad environmental gradients. Abiotic heterogeneity can result in a species that is the dominant competitor over some of its range being excluded if the area where it is dominant is too small. Biotic heterogeneity can lead to alternative stable landscapes or regional priority effects, while abiotic heterogeneity results in regional determinism. Broad environmental gradients in resource supply favor regional coexistence of species that exhibit local ASE.     

Eco-evolutionary feedbacks in community and ecosystem ecology: interactions between the ecological theatre and the evolutionary play

Interactions between natural selection and environmental change are well recognized and sit at the core of ecology and evolutionary biology. Reciprocal interactions between ecology and evolution, eco-evolutionary feedbacks, are less well studied, even though they may be critical for understanding the evolution of biological diversity, the structure of communities and the function of ecosystems. Eco-evolutionary feedbacks require that populations alter their environment (niche construction) and that those changes in the environment feed back to influence the subsequent evolution of the population. There is strong evidence that organisms influence their environment through predation, nutrient excretion and habitat modification, and that populations evolve in response to changes in their environment at time-scales congruent with ecological change (contemporary evolution). Here, we outline how the niche construction and contemporary evolution interact to alter the direction of evolution and the structure and function of communities and ecosystems. We then present five empirical systems that highlight important characteristics of eco-evolutionary feedbacks: rotifer–algae chemostats; alewife–zooplankton interactions in lakes; guppy life-history evolution and nutrient cycling in streams; avian seed predators and plants; and tree leaf chemistry and soil processes. The alewife–zooplankton system provides the most complete evidence for eco-evolutionary feedbacks, but other systems highlight the potential for eco-evolutionary feedbacks in a wide variety of natural systems.     

How understanding aboveground-belowground linkages can assist restoration ecology

The topic of aboveground-belowground linkages has seen much recent activity, resulting in several conceptual advances regarding plant-soil feedbacks, multitrophic interactions, and how organisms drive ecosystem processes. Although restoration ecology has been rapidly evolving as a scientific discipline, the principles that have developed regarding aboveground-belowground linkages have yet to be thoroughly integrated into it. In this review, we conceptually integrate the role of aboveground-belowground linkages with the principles of restoration ecology through a framework that transcends multiple levels of ecological organization, and illustrate its application through three examples: restoration of abandoned land, reversal of biological invasions, and restoration of natural disturbances. We conclude that this integration can greatly assist restoration ecology, through aiding identification of effective invention practices and prediction of ecosystem recovery.     

Alternative stable states in communities with intraguild predation

Intraguild predation (IGP) is a widespread ecological phenomenon in which two consumers that share a resource also engage in a predator-prey interaction. Theory on IGP predicts the occurrence of alternative stable states, but empirical evidence of such states is scarce. This raises the question of whether alternative states are a rare phenomenon that is unlikely to be observed in nature. Here we analyze a model in which the resource exhibits logistic or chemostat dynamics and consumers have saturating (Type II) functional responses. We show that alternative states can arise for a wide range of biological scenarios and that environmental constraints can make their detection difficult. Our analysis identifies three possible combinations of alternative states: (i) IG prey or IG predator, (ii) coexistence or IG predator, and (iii) coexistence or IG prey. Bifurcation diagrams reveal that alternative states are possible over large regions of the parameter space. However, they can be limited to narrow ranges along the resource productivity axis, which may make it difficult to observe the occurrence of alternative states in communities with IGP. Microcosm experiments provide a promising avenue for detecting combinations of asymptotically stable states along a productivity gradient.     

Phylogenetic relatedness as a tool in restoration ecology: a meta-analysis

Biotic interactions assembling plant communities can be positive (facilitation) or negative (competition) and operate simultaneously. Facilitative interactions and posterior competition are among the mechanisms triggering succession, thus representing a good scenario for ecological restoration. As distantly related species tend to have different phenotypes, and therefore different ecological requirements, they can coexist, maximizing facilitation and minimizing competition. We suggest including phylogenetic relatedness together with phenotypic information as a predictor for the net effects of the balance between facilitation and competition in nurse-based restoration experiments. We quantify, by means of a Bayesian meta-analysis of nurse-based restoration experiments performed worldwide, the importance of phylogenetic relatedness and life-form disparity in the survival, growth and density of facilitated plants. We find that the more similar the life forms of neighbouring plants are the greater the positive effect of phylogenetic distance is on survival and density. This result suggests that other characteristics beyond life form are also contained in the phylogeny, and the larger the phylogenetic distance, the less is the niche overlap, and therefore the less is the competition. As a general rule, we can maximize the success of the nurse-based practices by increasing life-form disparity and phylogenetic distances between the neighbour and the facilitated plant.     

Germination and emergence of annual species and burial depth: Implications for restoration ecology

Due to the high content of viable seeds, topsoil is usually spread on ground left bare during railway and motorway construction to facilitate the regeneration of vegetation cover. However, during handling of the topsoil, seeds are often buried deeply and they cannot germinate or the seedlings cannot emerge from depth. This study experimentally explores the predictive value of seed mass for seed germination, mortality and seedling emergence at different burial depths for 13 common annual species in semiarid Mediterranean environments. We separate the effect of burial depth on germination and emergence by means of two experiments. In the germination experiment, five replicates of 20 seeds for each species were buried at depths ranging from 0 to 4 cm under greenhouse conditions. Germinated and empty or rotten seeds were counted after 8 weeks. In the emergence experiment, five replicates of four newly-germinated seeds per species were buried at the same depths under controlled conditions and emergence was recorded after 3 weeks. The effect of burial depth on percentage of germination and seedling emergence was dependent on seed size. Although all species showed a decrease in germination with burial depth, this decrease was greater for small-than large-seeded species. Percentage of emergence was positively related to seed mass but negatively related to burial depth. Seed mortality was higher for small-than large-seeded species, but there was no general effect of burial depth on this variable. Thus, the current practice of spreading 30 cm deep layers of topsoil in post-construction restoration projects is unadvisable. In this restoration scenario, thinner layers of topsoil should be used to achieve the maximum potential of the topsoil for germination and seedling establishment.