Successional dynamics and alternative stable states in a saline activated sludge microbial community over 9 years

Background Microbial communities in both natural and applied settings reliably carry out myriads of functions, yet how stable these taxonomically diverse assemblages can be and what causes them to transition between states remains poorly understood. We studied monthly activated sludge (AS) samples collected over 9 years from a full-scale wastewater treatment plant to answer how complex AS communities evolve in the long term and how the community functions change when there is a disturbance in operational parameters. Results Here, we show that a microbial community in activated sludge (AS) system fluctuated around a stable average for 3 years but was then abruptly pushed into an alternative stable state by a simple transient disturbance (bleaching). While the taxonomic composition rapidly turned into a new state following the disturbance, the metabolic profile of the community and system performance remained remarkably stable. A total of 920 metagenome-assembled genomes (MAGs), representing approximately 70% of the community in the studied AS ecosystem, were recovered from the 97 monthly AS metagenomes. Comparative genomic analysis revealed an increased ability to aggregate in the cohorts of MAGs with correlated dynamics that are dominant after the bleaching event. Fine-scale analysis of dynamics also revealed cohorts that dominated during different periods and showed successional dynamics on seasonal and longer time scales due to temperature fluctuation and gradual changes in mean residence time in the reactor, respectively. Conclusions Our work highlights that communities can assume different stable states under highly similar environmental conditions and that a specific disturbance threshold may lead to a rapid shift in community composition.

Large herbivores suppress liana infestation in an African savanna

African savannas are the last stronghold of diverse large-mammal communities, and a major focus of savanna ecology is to understand how these animals affect the relative abundance of trees and grasses. However, savannas support diverse plant life-forms, and human-induced changes in large-herbivore assemblages—declining wildlife populations and their displacement by livestock—may cause unexpected shifts in plant community composition. We investigated how herbivory affects the prevalence of lianas (woody vines) and their impact on trees in an East African savanna. Although scarce (<2% of tree canopy area) and defended by toxic latex, the dominant liana, Cynanchum viminale (Apocynaceae), was eaten by 15 wild large-herbivore species and was consumed in bulk by native browsers during experimental cafeteria trials. In contrast, domesticated ungulates rarely ate lianas. When we experimentally excluded all large herbivores for periods of 8 to 17 y (simulating extirpation), liana abundance increased dramatically, with up to 75% of trees infested. Piecewise exclusion of different-sized herbivores revealed functional complementarity among size classes in suppressing lianas. Liana infestation reduced tree growth and reproduction, but herbivores quickly cleared lianas from trees after the removal of 18-y-old exclosure fences (simulating rewilding). A simple model of liana contagion showed that, without herbivores, the long-term equilibrium could be either endemic (liana–tree coexistence) or an all-liana alternative stable state. We conclude that ongoing declines of wild large-herbivore populations will disrupt the structure and functioning of many African savannas in ways that have received little attention and that may not be mitigated by replacing wildlife with livestock.

Fast environmental change and eco-evolutionary feedbacks can drive regime shifts in ecosystems before tipping points are crossed

Anthropogenic environmental changes are altering ecological and evolutionary processes of ecosystems. The possibility that ecosystems can respond abruptly to gradual environmental change when critical thresholds are crossed (i.e. tipping points) and shift to an alternative stable state is a growing concern. Here I show that fast environmental change can trigger regime shifts before environmental stress exceeds a tipping point in evolving ecological systems. The difference in the time scales of coupled ecological and evolutionary processes makes ecosystems sensitive not only to the magnitude of environmental changes, but also to the rate at which changes are imposed. Fast evolutionary change mediated by high trait variation can reduce the sensitivity of ecosystems to the rate of environmental change and prevent the occurrence of rate-induced regime shifts. This suggests that management measures to prevent rate-induced regime shifts should focus on mitigating the effects of environmental change and protecting phenotypic diversity in ecosystems.

Anticipating Critical Transitions in Psychological Systems using Early Warning Signals: Theoretical and Practical Considerations

Many real-world systems can exhibit sudden shifts from one stable state to another, and the theory of dynamical systems points to the existence of generic early warning signals that precede such shifts. Recently, psychologists have begun to conceptualize mental disorders such as depression as an alternative stable state, and suggested that early warning signals based on the phenomenon of critical slowing down might be useful for predicting sudden transitions into depression or other psychiatric disorders. Harnessing the potential of early warning signals requires us to understand their limitations as well as the factors influencing their performance in practice. In this paper, we (a) review limitations of early warning signals based on critical slowing down to better understand when they can and cannot occur, and (b) study the conditions under which early warning signals may anticipate critical transitions in online-monitoring settings by simulating from a bistable dynamical system, varying crucial features such as sampling frequency, noise intensity, and speed of approaching the tipping point. We find that, in sharp contrast to their reputation of being generic or model-agnostic, whether early warning signals occur or not strongly depends on the specifics of the system. We also find that they are very sensitive to noise, potentially limiting their utility in practical applications. We discuss the implications of our findings and provide suggestions and recommendations for future research.

Biocrust as one of multiple stable states in global drylands

Biocrusts cover ~30% of global drylands with a prominent role in the biogeochemical cycles. Theoretically, biocrusts, vascular plants, and bare soil can represent multiple stable states in drylands. However, no empirical evidence for the existence of a biocrust stable state has been reported. Here, using a global drylands dataset, we found that biocrusts form an alternative stable state (biocrust cover, ~80%; vascular cover, ≤10%) besides bare soil (both biocrust and vascular cover, ≤10%) and vascular plants (vascular cover, >50%; biocrust cover, ~5%). The pattern of multiple stable states associated with biocrusts differs from the classic fold bifurcation, and values of the aridity index in the range of 0 to 0.6 define a bistable region where multiple stable states coexist. This study empirically demonstrates the existence and thresholds of multiple stable states associated with biocrusts along climatic gradients and thus may greatly contribute to conservation and restoration of global drylands.

Different coastal marsh sites reflect similar topographic conditions for bare patches and vegetation recovery

The presence of bare patches within otherwise vegetated coastal marshes is sometimes considered to be a symptom of marsh die-back and the subsequent loss of important ecosystem services. Here we studied the topographical conditions determining the presence and revegetation of bare patches in three marsh sites with contrasting tidal range, sediment supply and plant species: the Scheldt Estuary (the Netherlands), Venice Lagoon (Italy), and Blackwater Marshes (Maryland, USA). We analyzed topographic properties of bare patches, including elevation, size, distance and connectivity to channels based on GIS analyses of aerial and LIDAR imagery. Our results demonstrate that across the different marsh sites, bare patches connected to channels occur most frequently at the lowest elevations and farthest distance from the main channels. Bare patches disconnected from channels occur most frequently at intermediate elevations and distances from channels, and vegetated marshes dominate at highest elevations and shortest distances from channels. Revegetation in bare patches is observed in only one site with the highest tidal range and highest sediment availability, and preferentially occurs from the edges of small unconnected bare patches at intermediate elevations and intermediate distances from channels. Our results are discussed within the alternative stable state theory. We suggest the existence of two stable states, a high-elevated vegetated state close to channels that tends to remain high and vegetated, and a low-elevated state of bare connected patches far from channels that tends to remain bare, with an unstable state at intermediate channel distances where bare patches may form and rapidly become revegetated.

Transient invaders can induce shifts between alternative stable states of microbial communities

Microbial dispersal often leads to the arrival of outsider organisms into ecosystems. When their arrival gives rise to successful invasions, outsider species establish within the resident community, which can markedly alter the ecosystem. Seemingly less influential, the potential impact of unsuccessful invaders that interact only transiently with the community has remained largely ignored. Here, we experimentally demonstrate that these transient invasions can induce a lasting transition to an alternative stable state, even when the invader species itself does not survive the transition. First, we develop a mechanistic understanding of how environmental changes caused by these transient invaders can drive a community shift in a simple, bistable model system. Beyond this, we show that transient invaders can also induce switches between stable states in more complex communities isolated from natural soil samples. Our results demonstrate that short-term interactions with an invader species can induce lasting shifts in community composition and function.

A palaeolimnological perspective to understand regime-shift dynamics in two Yangtze-basin lakes

Natural and human disturbances have caused widespread regime shifts in shallow lakes of the lower Yangtze basin (LYB, China) resulting in a severe decline of ecosystem services. Improved understanding of the relationship between environmental forcing and ecosystem response, and the mechanisms behind regime shifts has significant implications for management. However, the patterns of these regime shifts and the underlying internal mechanisms are less known. In this study, two typical lakes (Chaohu and Zhangdu) from the LYB were selected to determine the trajectories of ecological regime shifts, both of which transitioned from vegetation- to plankton-dominated states several decades ago. Ecological trajectories since the 1900s in both lakes were reconstructed using palaeolimnological proxies, mainly diatom assemblages. Although results show that regime shifts occurred in both lakes in the 1970s and the 1950s, respectively, their inherent mechanisms were different. In Lake Zhangdu, altered hydrological conditions pushed the ecosystem across an ecological threshold, providing an example of a driver-mediated regime shift. In Lake Chaohu, ongoing nutrient loading influenced ecosystem processes and drove the lake to an alternative stable state, potentially presenting an example of a critical transition after a loss of resilience. This research indicates that palaeolimnological perspectives can provide insights into regime shift changes, as well as important information regarding which restoration methods should be tailored to individual lakes.

Uncertain ecosystems: the conceptual framework

Climate sets the potential biomass of trees and physiologists have made considerable progress in understanding and predicting that potential and applying it in global vegetation models. The problem is in understanding and predicting tree cover where it is far from the climate potential. Vast areas of non-forested vegetation occur where climates are suitable for forests. Arguments over why forests are absent, ongoing for over a century, are generally polarized between favouring bottom-up factors (resource constraints) or top-down factors (herbivory, predation, fire). There is increasing support for hypotheses invoking the interaction between the two. This chapter introduces the key hypotheses, their assumptions and predictions. Trophic ecology is a useful framework for exploring departures from the climate potential for trees, focussing explicitly on regulation by consumers, including fire. Alternative stable state theory is emerging as particularly appropriate for explaining forest/non-forest mosaics with each state maintained by positive feedbacks to the preferred environment.