Eco-evolutionary litter feedback as a driver of exotic plant invasion

Many studies have examined positive feedbacks between invasive plant traits and nutrient cycling, but few have investigated whether feedbacks arise from introduction of pre-adapted species or from eco-evolutionary feedback that develops after introduction. Eco-evolutionary feedback could occur between an invader's leaf tissue C:N ratio and its response to litter accumulation. Previous modeling predicts that occurrence of this feedback would be reflected by: (1) field data showing higher litter:biomass ratios in the invasive range; (2) high C:N genotypes benefiting more from experimental litter additions than low C:N genotypes; (3) this beneficial effect on high C:N genotypes inducing a critical transition toward invader dominance when a critical amount of litter is added to a native species-dominated community experiencing low nutrient conditions. Here, we empirically tested these predictions for the invasive grass Phalaris arundinacea, which has undergone post-introduction evolutionary change toward attaining higher C:N ratios under high nutrient conditions. We performed a biogeographical comparison of litter:biomass ratios in the native (Europe) and invasive (USA) range, and an experiment with mesocosms from the invasive range under low nutrient conditions. Low and high C:N Phalaris genotypes were introduced into native-dominated and bare mesocosms, to which varying litter amounts were added. The biogeographical comparison revealed that litter:biomass ratios were higher in the invasive range. The mesocosm experiment showed that when grown in isolation, only high C:N genotypes responded positively to litter. This effect, however, was not strong enough to stimulate Phalaris when exposed to competition with native species. Our results suggest that eco-evolutionary feedback between Phalaris’ C:N ratio and litter accumulation could occur, but only under high nutrient conditions. Our experiments suggest that eco-evolutionary feedback may select for specialist rather than superior genotypes. Hence, genotypic variation induced by post-introduction admixture may be subject to context-dependent selection due to eco-evolutionary feedback, increasing trait variation within invasive populations.     

Accounting for the nested nature of genetic variation across levels of organization improves our understanding of biodiversity and community ecology

Recent work has demonstrated that the presence or abundance of specific genotypes, populations, species and phylogenetic clades may influence community and ecosystem properties such as resilience or productivity. Many ecological studies, however, use simple linear models to test for such relationships, including species identity as the predictor variable and some measured trait or function as the response variable without accounting for the nestedness of genetic variation across levels of organization. This omission may lead to incorrect inference about which source of variation influences community and ecosystem properties. Here, we explicitly compare this common approach to alternative ways of modeling variation in trait data, using simulated trait data and empirical results of common‐garden trials using multiple levels of genetic variation within Eucalyptus, Populus and Picea. We show that: 1) when nested variation is ignored, an incorrect conclusion of species effect is drawn in up to 20% of cases; 2) overestimation of the species effect increases – up to 60% in some scenarios – as the nested term explains more of the variation; and 3) the sample sizes needed to overcome these potential problems associated with aggregating nested hierarchical variation may be impractically large. In common‐garden trials, incorporating nested models increased explanatory power twofold for mammal browsing rate in Eucalyptus, threefold for leaf area in Populus, and tenfold for branch number in Picea. Thoroughly measuring intraspecific variation and characterizing hierarchical genetic variation beyond the species level has implications for developing more robust theory in community ecology, managing invaded natural systems, and improving inference in biodiversity–ecosystem functioning research. Synthesis Until recently, ecologists acknowledged the ubiquity of within‐species trait variation, but paid scant attention to how much it affects communities and ecosystems. Here, the authors used simulated trait data and common‐garden studies to demonstrate that we ignore intraspecific trait variation at our peril. In both simulated and experimental systems, in many cases ignoring intraspecific variation led to incorrect statistical inferences and inflated the effect size of species identity. This study shows that ecologists must characterize hierarchically nested genetic and phenotypic variation to fully understand the links between individual traits, community structure and ecosystem functioning.     

When and where plant‐soil feedback may promote plant coexistence: a meta‐analysis

Plant‐soil feedback (PSF) theory provides a powerful framework for understanding plant dynamics by integrating growth assays into predictions of whether soil communities stabilise plant–plant interactions. However, we lack a comprehensive view of the likelihood of feedback‐driven coexistence, partly because of a failure to analyse pairwise PSF, the metric directly linked to plant species coexistence. Here, we determine the relative importance of plant evolutionary history, traits, and environmental factors for coexistence through PSF using a meta‐analysis of 1038 pairwise PSF measures. Consistent with eco‐evolutionary predictions, feedback is more likely to mediate coexistence for pairs of plant species (1) associating with similar guilds of mycorrhizal fungi, (2) of increasing phylogenetic distance, and (3) interacting with native microbes. We also found evidence for a primary role of pathogens in feedback‐mediated coexistence. By combining results over several independent studies, our results confirm that PSF may play a key role in plant species coexistence, species invasion, and the phylogenetic diversification of plant communities.     

Tropomyosin and caldesmon regulate cytokinesis speed and membrane stability during cell division

The contractile ring and the cell cortex generate force to divide the cell while maintaining symmetrical shape. This requires temporal and spatial regulation of the actin cytoskeleton at these areas. We force-expressed misregulated versions of actin-binding proteins, tropomyosin and caldesmon, into cells and analyzed their effects on cell division. Cells expressing proteins that increase actomyosin ATPase, such as human tropomyosin chimera (hTM5/3), significantly speed up division, whereas cells expressing proteins that inhibit actomyosin, such as caldesmon mutants defective in Ca(2+)/calmodulin binding (CaD39-AB) and in cdk1 phosphorylation sites (CaD39-6F), divide slowly. hTM5 and hTM5/3-expressing cells lift one daughter cell off the substrate and twist. Furthermore, CaD39-AB- and CaD39-6F-expressing cells are sensitive to hypotonic swelling and show severe blebbing during division, whereas hTM5/3-expressing cells are resistant to hypotonic swelling and produce membrane bulges. These results support a model where Ca(2+)/calmodulin and cdk1 dynamically control caldesmon inhibition of tropomyosin-activated actomyosin to regulate division speed and to suppress membrane blebs.     

Antigonon leptopus invasion is associated with plant community disassembly in a Caribbean island ecosystem

Biological Invasions - Invasions by non-native plant species are widely recognized as a major driver of biodiversity loss. Globally, (sub-)tropical islands form important components of biodiversity...     

The effect of climate change on the resilience of ecosystems with adaptive spatial pattern formation

In a rapidly changing world, quantifying ecosystem resilience is an important challenge. Historically, resilience has been defined via models that do not take spatial effects into account. These systems can only adapt via uniform adjustments. In reality, however, the response is not necessarily uniform, and can lead to the formation of (self‐organised) spatial patterns – typically localised vegetation patches. Classical measures of resilience cannot capture the emerging dynamics in spatially self‐organised systems, including transitions between patterned states that have limited impact on ecosystem structure and productivity. We present a framework of interlinked phase portraits that appropriately quantifies the resilience of patterned states, which depends on the number of patches, the distances between them and environmental conditions. We show how classical resilience concepts fail to distinguish between small and large pattern transitions, and find that the variance in interpatch distances provides a suitable indicator for the type of imminent transition. Subsequently, we describe the dependency of ecosystem degradation based on the rate of climatic change: slow change leads to sporadic, large transitions, whereas fast change causes a rapid sequence of smaller transitions. Finally, we discuss how pre‐emptive removal of patches can minimise productivity losses during pattern transitions, constituting a viable conservation strategy.     

Plant–soil feedbacks and the coexistence of competing plants

Plant–soil feedbacks can have important implications for the interactions among plants. Understanding these effects is a major challenge since it is inherently difficult to measure and manipulate highly diverse soil communities. Mathematical models may advance this understanding by making the interplay of the various processes affecting plant–soil interaction explicit and by quantifying the relative importance of the factors involved. The aim of this paper is to provide a complete analysis of a pioneering plant–soil feedback model developed by Bever and colleagues (J Ecol 85: 561–573, 1997; Ecol Lett 2: 52–62, 1999; New Phytol 157: 465–473, 2003) to fully understand the range of possible impacts of plant–soil feedbacks on plant communities within this framework. We analyze this model by means of a new graphical method that provides a complete classification of the potential effects of soil communities on plant competition. Due to the graphical character of the method, the results are relatively easy to obtain and understand. We show that plant diversity depends crucially on two key parameters that may be viewed as measures of the intensity of plant competition and the direction and strength of plant–soil feedback, respectively. Our analysis provides a formal underpinning of earlier claims that plant–soil feedbacks, especially when they are negative, may enhance the diversity of plant communities. In particular, negative plant–soil feedbacks can enhance the range of plant coexistence by inducing competitive oscillations. However, these oscillations can also destabilize plant coexistence, leading to low population densities and extinctions. In addition, positive feedbacks can allow locally stable forms of plant coexistence by inducing alternative stable states. Our findings highlight that the inclusion of plant–soil interactions may fundamentally alter the predictions on the structure and functioning of above-ground ecosystems. The scenarios presented in this study can be used to formulate hypotheses about the ways soil community effects may influence plant competition that can be tested with empirical studies. This will advance our understanding of the role of plant–soil feedback in ecological communities.     

Opposite effects of overexpressed myosin Va or heavy meromyosin Va on vesicle distribution, cytoskeleton organization, and cell motility in nonmuscle cells

Myosin Va, an actin-based motor protein that transports intracellular cargos, can bundle actin in vitro. Whether myosin Va regulates cellular actin dynamics or cell migration remains unclear. To address this, we compared Chinese Hamster Ovary (CHO) cells that stably express GFP fused to either full length mouse myosin Va (GFP-M5) or heavy meromyosin Va (GFP-M5Delta). GFP-M5 and GFP-M5Delta co-immunoprecipitate with CHO myosin Va and serve as overexpression of wild-type and dominant negative mutants of myosin Va. Compared to non-expressing control cells, GFP-M5-overexpressing cells have peripheral endocytic vesicles, spread slowly after plating, as well as produce robust interior actin stress fibers, myosin II bundles, and focal adhesions. However, these cells display normal cell migration and lamellipodial dynamics. In contrast, GFP-M5Delta-expressing cells have perinuclear endocytic vesicles, produce thin interior actin and myosin bundles and contain no interior focal adhesions. In addition, these cells spread rapidly, migrate slowly and display reduced lamellipodial dynamics. Similarly, neurite outgrowth is compromised in neurons cultured from transgenic Drosophila that express M5Delta-dsRed and in neurons cultured from Drosophila that produce a tailless version of endogenous myosin V. Together, these data suggest that myosin Va overexpression induces actin bundles in vivo whereas the tailless version fails to bundle actin and disrupts cell motility.     

Ecosystems off track: rate‐induced critical transitions in ecological models

Theory suggests that gradual environmental change may erode the resilience of ecosystems and increase their susceptibility to critical transitions. This notion has received a lot of attention in ecology in recent decades. An important question receiving far less attention is whether ecosystems can cope with the rapid environmental changes currently imposed. The importance of this question was recently highlighted by model studies showing that elevated rates of change may trigger critical transitions, whereas slow environmental change would not. This paper aims to provide a mechanistic understanding of these rate‐induced critical transitions to facilitate identification of rate sensitive ecosystems. Analysis of rate sensitive ecological models is challenging, but we demonstrate how rate‐induced transitions in an elementary model can still be understood. Our analyses reveal that rate‐induced transitions 1) occur if the rate of environmental change is high compared to the response rate of ecosystems, 2) are driven by rates, rather than magnitudes, of change and 3) occur once a critical rate of change is exceeded. Disentangling rate‐induced transitions from classical transitions in observations would be challenging. However, common features of rate‐sensitive models suggest that ecosystems with coupled fast–slow dynamics, exhibiting repetitive catastrophic shifts or displaying periodic spatial patterns are more likely to be rate sensitive. Our findings are supported by experimental studies showing rate‐dependent outcomes. Rate sensitivity of models suggests that the common definition of ecological resilience is not suitable for a subset of real ecosystems and that formulating limits to magnitudes of change may not always safeguard against ecosystem degradation. Synthesis Understanding and predicting ecosystem response to environmental change is one of the key challenges in ecology. Model studies have suggested that slow, gradual environmental change beyond some critical threshold can trigger so‐called critical transitions and abrupt ecosystem degradation. An important question remains however whether ecosystems can cope with the ongoing rapid anthropogenic environmental changes to which they are currently imposed. In this study we demonstrate that in some ecological models elevated rates of change can trigger critical transitions even if slow environmental change of the same magnitude would not. Such rateinduced critical transitions in models suggest that concepts like resilience and planetary boundaries may not always be sufficient to explain and prevent ecosystem degradation.     

Empirical and theoretical challenges in aboveground–belowground ecology

A growing body of evidence shows that aboveground and belowground communities and processes are intrinsically linked, and that feedbacks between these subsystems have important implications for community structure and ecosystem functioning. Almost all studies on this topic have been carried out from an empirical perspective and in specific ecological settings or contexts. Belowground interactions operate at different spatial and temporal scales. Due to the relatively low mobility and high survival of organisms in the soil, plants have longer lasting legacy effects belowground than aboveground. Our current challenge is to understand how aboveground–belowground biotic interactions operate across spatial and temporal scales, and how they depend on, as well as influence, the abiotic environment. Because empirical capacities are too limited to explore all possible combinations of interactions and environmental settings, we explore where and how they can be supported by theoretical approaches to develop testable predictions and to generalise empirical results. We review four key areas where a combined aboveground–belowground approach offers perspectives for enhancing ecological understanding, namely succession, agro-ecosystems, biological invasions and global change impacts on ecosystems. In plant succession, differences in scales between aboveground and belowground biota, as well as between species interactions and ecosystem processes, have important implications for the rate and direction of community change. Aboveground as well as belowground interactions either enhance or reduce rates of plant species replacement. Moreover, the outcomes of the interactions depend on abiotic conditions and plant life history characteristics, which may vary with successional position. We exemplify where translation of the current conceptual succession models into more predictive models can help targeting empirical studies and generalising their results. Then, we discuss how understanding succession may help to enhance managing arable crops, grasslands and invasive plants, as well as provide insights into the effects of global change on community re-organisation and ecosystem processes.