The temporal and spatial dimensions of plant-soil feedbacks.

Feedbacks between plants and soil microbes form a keystone to terrestrial community and ecosystem dynamics. Recent advances in dissecting the spatial and temporal dynamics of plant-soil feedbacks (PSFs) have challenged longstanding assumptions of spatially well-mixed microbial communities and exceedingly fast microbial assembly dynamics relative to plant lifespans. Instead, PSFs emerge from interactions that are inherently mismatched in spatial and temporal scales, and explicitly considering these spatial and temporal dynamics is crucial to understanding the contribution of PSFs to foundational ecological patterns. I propose a synthetic spatiotemporal framework for future research that pairs experimental and modeling approaches grounded in mechanism to improve predictability and generalizability of PSFs.

Improving Instructional Fitness Requires Change.

Transmission of information has benefitted from a breathtaking level of innovation and change over the past 20 years; however, instructional methods within colleges and universities have been slow to change. In the article, we present a novel framework to structure conversations that encourage innovation, change, and improvement in our system of higher education, in general, and our system of biology education, specifically. In particular, we propose that a conceptual model based on evolutionary landscapes in which fitness is replaced by educational effectiveness would encourage educational improvement by helping to visualize the multidimensional nature of education and learning, acknowledge the complexity and dynamism of the educational landscape, encourage collaboration, and stimulate experimental thinking about how new approaches and methodology could take various fields associated with learning, to more universal fitness optima. The framework also would encourage development and implementation of new techniques and persistence through less efficient or effective valleys of death.

Divergence in Diversity and Composition of Root-Associated Fungi Between Greenhouse and Field Studies in a Semiarid Grassland.

Investigations of plant-soil feedbacks (PSF) and plant-microbe interactions often rely exclusively on greenhouse experiments, yet we have little understanding of how, and when, results can be extrapolated to explain phenomena in nature. A systematic comparison of microbial communities using the same host species across study environments can inform the generalizability of such experiments. We used Illumina MiSeq sequencing to characterize the root-associated fungi of two foundation grasses from a greenhouse PSF experiment, a field PSF experiment, field monoculture stands, and naturally occurring resident plants in the field. A core community consisting < 10% of total fungal OTU richness but > 50% of total sequence abundance occurred in plants from all study types, demonstrating the ability of field and greenhouse experiments to capture the dominant component of natural communities. Fungal communities were plant species-specific across the study types, with the core community showing stronger host specificity than peripheral taxa. Roots from the greenhouse and field PSF experiments had lower among sample variability in community composition and higher diversity than those from naturally occurring, or planted monoculture plants from the field. Core and total fungal composition differed substantially across study types, and dissimilarity between fungal communities did not predict plant-soil feedbacks measured in experiments. These results suggest that rhizobiome assembly mechanisms in nature differ from the dynamics of short-term, inoculation studies. Our results validate the efficacy of common PSF experiment designs to test soil inoculum effects, and highlight the challenges of scaling the underlying microbial mechanisms of plant responses from whole-community inoculation experiments to natural ecosystems.

Climate sensitivity functions and net primary production: A framework for incorporating climate mean and variability.

Understanding controls on net primary production (NPP) has been a long-standing goal in ecology. Climate is a well-known control on NPP, although the temporal differences among years within a site are often weaker than the spatial pattern of differences across sites. Climate sensitivity functions describe the relationship between an ecological response (e.g., NPP) and both the mean and variance of its climate driver (e.g., aridity index), providing a novel framework for understanding how climate trends in both mean and variance vary with NPP over time. Nonlinearities in these functions predict whether an increase in climate variance will have a positive effect (convex nonlinearity) or negative effect (concave nonlinearity) on NPP. The influence of climate variance may be particularly intense at ecosystem transition zones, if species reach physiological thresholds that create nonlinearities at these ecotones. Long-term data collected at the confluence of three dryland ecosystems in central New Mexico revealed that each ecosystem exhibited a unique climate sensitivity function that was consistent with long-term vegetation change occurring at their ecotones. Our analysis suggests that rising temperatures in drylands could alter the nonlinearities that determine the relative costs and benefits of variance in precipitation for primary production.

Plant-soil feedbacks promote negative frequency dependence in the coexistence of two aridland grasses.

Understanding the mechanisms of species coexistence is key to predicting patterns of species diversity. Historically, the ecological paradigm has been that species coexist by partitioning resources: as a species increases in abundance, self-limitation kicks in, because species-specific resources decline. However, determining coexistence mechanisms has been a particular puzzle for sedentary organisms with high overlap in their resource requirements, such as plants. Recent evidence suggests that plant-associated microbes could generate the stabilizing self-limitation (negative frequency dependence) that is required for species coexistence. Here, we test the key assumption that plant-microbe feedbacks cause such self-limitation. We used competition experiments and modelling to evaluate how two common groups of soil microbes (rhizospheric microbes and biological soil crusts) influenced the self-limitation of two competing desert grass species. Negative feedbacks between the dominant plant competitor and its rhizospheric microbes magnified self-limitation, whereas beneficial interactions between both plant species and biological soil crusts partly counteracted this stabilizing effect. Plant-microbe interactions have received relatively little attention as drivers of vegetation dynamics in dry land ecosystems. Our results suggest that microbial mechanisms can contribute to patterns of plant coexistence in arid grasslands.

Greater sexual reproduction contributes to differences in demography of invasive plants and their noninvasive relatives.

An understanding of the demographic processes contributing to invasions would improve our mechanistic understanding of the invasion process and improve the efficiency of prevention and control efforts. However, field comparisons of the demography of invasive and noninvasive species have not previously been conducted. We compared the in situ demography of 17 introduced plant species in St. Louis, Missouri, USA, to contrast the demographic patterns of invasive species with their less invasive relatives across a broad sample of angiosperms. Using herbarium records to estimate spread rates, we found higher maximum spread rates in the landscape for species classified a priori as invasive than for noninvasive introduced species, suggesting that expert classifications are an accurate reflection of invasion rate. Across 17 species, projected population growth was not significantly greater in invasive than in noninvasive introduced species. Among five taxonomic pairs of close relatives, however, four of the invasive species had higher projected population growth rates compared with their noninvasive relative. A Life Table Response Experiment suggested that the greater projected population growth rate of some invasive species relative to their noninvasive relatives was primarily a result of sexual reproduction. The greater sexual reproduction of invasive species is consistent with invaders having a life history strategy more reliant on fecundity than survival and is consistent with a large role of propagule pressure in invasion. Sexual reproduction is a key demographic correlate of invasiveness, suggesting that local processes influencing sexual reproduction, such as enemy escape, might be of general importance. However, the weak correlation of projected population growth with spread rates in the landscape suggests that regional processes, such as dispersal, may be equally important in determining invasion rate.

Do plant-soil feedbacks promote coexistence in a sagebrush steppe?

Recent studies have shown the potential for negative plant-soil feedbacks (PSFs) to promote stable coexistence, but have not quantified the stabilizing effect relative to other coexistence mechanisms. We conducted a field experiment to test the role of PSFs in stabilizing coexistence among four dominant sagebrush steppe species that appear to coexist stably, based on previous work with observational data and models. We then integrated the effects of PSF treatments on focal species across germination, survival, and first-year growth. To contribute to stable coexistence, soil microbes should have host-specific effects that result in negative feedbacks. Over two replicated growing seasons, our experiments consistently showed that soil microbes have negative effects on plant growth, but these effects were rarely host-specific. The uncommon host-specific effects were mostly positive at the germination stage, and negative for growth. Integrated effects of PSF across early life-stage vital rates showed that PSF-mediated self-limitation occasionally had large effects on projected plant biomass, but occurred inconsistently between years. Our results suggest that while microbially-mediated PSF may not be a common mechanism of coexistence in this community, it may still affect the relative abundance of dominant plant species via changes in host fitness. Our work also serves as a blueprint for future investigations that aim to identify underlying processes and test alternative mechanisms to explain important patterns in community ecology.

Connecting plant-soil feedbacks to long-term stability in a desert grassland.

Temporal fluctuations in plant species coexistence are key to understanding ecosystem state transitions and long-term maintenance of species diversity. Although plant microbiomes can alter plant competition in short-term experiments, their relevance to natural temporal patterns in plant communities is unresolved. In a semiarid grassland, the frequency and magnitude of change in plant species composition through time varied from relatively static to highly dynamic among patches across the landscape. We field tested whether these alternative successional trajectories correlated with alternative plant-soil interactions. In temporally stable patches, we found negative plant-soil feedbacks, where plants grew worse with conspecific than heterospecific soil biota-a mechanism that maintains stability in mathematical models. In contrast, feedbacks in temporally dynamic patches were neutral to positive. Importantly, the magnitude of feedbacks depended on plant frequency, enabling plant species to increase in cover when rare, which theory predicts will promote long-term, stable coexistence. Although our study does not determine the direction of causality, our results reveal a novel link between plant-microbe interactions and temporal stability of plant species coexistence and help to explain 20+ yr of plant abundance dynamics at the patch-to-landscape scales.

A mutualistic endophyte alters the niche dimensions of its host plant.

Mutualisms can play important roles in influencing species coexistence and determining community composition. However, few studies have tested whether such interactions can affect species distributions by altering the niches of partner species. In subalpine meadows of the Rocky Mountains, USA, we explored whether the presence of a fungal endophyte (genus Epichloë) may shift the niche of its partner plant, marsh bluegrass (Poa leptocoma) relative to a closely related but endophyte-free grass species, nodding bluegrass (Poa reflexa). Using observations and a 3-year field experiment, we tested two questions: (i) Do P. leptocoma and P. reflexa occupy different ecological niches? and (ii) Does endophyte presence affect the relative fitness of P. leptocoma versus P. reflexa in the putative niches of these grass species? The two species were less likely to co-occur than expected by chance. Specifically, P. leptocoma grew closer to water sources and in wetter soils than P. reflexa, and also had higher root colonization by mycorrhizal fungi. Endophyte-symbiotic P. leptocoma seeds germinated with greater frequency in P. leptocoma niches relative to P. reflexa niches, whereas neither endophyte-free (experimentally removed) P. leptocoma seeds nor P. reflexa seeds showed differential germination between the two niche types. Thus, endophyte presence constrained the germination and early survival of host plants to microsites occupied by P. leptocoma. However, endophyte-symbiotic P. leptocoma ultimately showed greater growth than endophyte-free plants across all microsites, indicating a net benefit of the symbiosis at this life history stage. Differential effects of endophyte symbiosis on different host life history stages may thus contribute to niche partitioning between the two congeneric plant species. Our study therefore identifies a symbiotic relationship as a potential mechanism facilitating the coexistence of two species, suggesting that symbiont effects on host niche may have community-level consequences.

Minimal effects of an invasive flowering shrub on the pollinator community of native forbs.

Biological invasions can strongly influence species interactions such as pollination. Most of the documented effects of exotic plant species on plant-pollinator interactions have been observational studies using single pairs of native and exotic plants, and have focused on dominant exotic plant species. We know little about how exotic plants alter interactions in entire communities of plants and pollinators, especially at low to medium invader densities. In this study, we began to address these gaps by experimentally removing the flowers of a showy invasive shrub, Rosa multiflora, and evaluating its effects on the frequency, richness, and composition of bee visitors to co-flowering native plants. We found that while R. multiflora increased plot-level richness of bee visitors to co-flowering native plant species at some sites, its presence had no significant effects on bee visitation rate, visitor richness, bee community composition, or abundance overall. In addition, we found that compared to co-flowering natives, R. multiflora was a generalist plant that primarily received visits from generalist bee species shared with native plant species. Our results suggest that exotic plants such as R. multiflora may facilitate native plant pollination in a community context by attracting a more diverse assemblage of pollinators, but have limited and idiosyncratic effects on the resident plant-pollinator network in general.