Global patterns and drivers of plant-soil microbe interactions.

Plant-soil feedback (PSF) is an important mechanism determining plant community dynamics and structure. Understanding the geographic patterns and drivers of PSF is essential for understanding the mechanisms underlying geographic plant diversity patterns. We compiled a large dataset containing 5969 observations of PSF from 202 studies to demonstrate the global patterns and drivers of PSF for woody and non-woody species. Overall, PSF was negative on average and was influenced by plant attributes and environmental settings. Woody species PSFs did not vary with latitude, but non-woody PSFs were more negative at higher latitudes. PSF was consistently more positive with increasing aridity for both woody and non-woody species, likely due to increased mutualistic microbes relative to soil-borne pathogens. These findings were consistent between field and greenhouse experiments, suggesting that PSF variation can be driven by soil legacies from climates. Our findings call for caution to use PSF as an explanation of the latitudinal diversity gradient and highlight that aridity can influence plant community dynamics and structure across broad scales through mediating plant-soil microbe interactions.

Responses of grassland productivity to mowing intensity and precipitation variability in a temperate steppe.

Mowing for hay is an important land use in grasslands that is affected by precipitation variability, due to the water-limited nature of these ecosystems. Past land use and precipitation conditions can have legacy effects on ecosystem functions, potentially altering responses to both mowing and precipitation. Nonetheless, it is still unclear how natural variation in precipitation will affect plant responses to changes in mowing intensity. We conducted a seven-year field experiment with three mowing intensity treatments compared to the traditional mowing intensity (5 cm stubble height) as a control: increased mowing (2 cm stubble), decreased mowing (8 cm stubble) and ceased mowing. Decreased mowing increased both plant aboveground net primary productivity [ANPP] and forage yield across the whole community, driven by increases in graminoids, mainly owing to the positive response of plants to precipitation. Both mowing disturbance and precipitation variability had legacy effects on plant ANPP; however, these responses differed among the whole community, graminoid, and forb levels. Current-year community-wide ANPP [ANPPn] was positively associated with current-year precipitation [PPTn] in all mowing treatments, driven by positive precipitation responses of the dominant graminoids. For forbs, however, ANPPn was negatively associated with prior-year growing season precipitation [PPTn-1] across mowing treatments, potentially due to lagged competition with the dominant graminoids. Our results suggest that the response of the dominant graminoids is the primary factor determining the response of ANPP to mowing and precipitation variability in these grassland ecosystems, and highlight that decreasing mowing intensity may maximize both herder's income and grassland sustainability.

Plant responses to soil biota depend on precipitation history, plant diversity, and productivity.

Soil biota are critical drivers of plant growth, population dynamics, and community structure and thus have wide-ranging effects on ecosystem function. Interactions between plants and soil biota are complex, however, and can depend on the diversity and productivity of the plant community and environmental conditions. Plant-soil biota interactions may be especially important during stressful periods, such as drought, when plants can gain great benefits from beneficial biota but may be susceptible to antagonists. How soil biota respond to drought is also important and can influence plant growth following drought and leave legacies that affect future plant responses to soil biota and further drought. To explore how drought legacies and plant community context influence plant growth responses to soil biota and further drought, we collected soils from 12 grasslands varying in plant diversity and productivity where precipitation was experimentally reduced. We used these soils as inoculum in a growth chamber experiment testing how precipitation history (ambient or reduced) and soil biota (live or sterile soil inoculum) mediate plant growth and drought responses within an experimental plant community. We also tested whether these responses differed with the diversity and productivity of the community where the soil was collected. Plant growth responses to soil biota were positive when inoculated with soils from less diverse and productive plant communities and became negative as the diversity and productivity of the conditioning community increased. At low diversity, however, positive soil biota effects on plant growth were eliminated if precipitation had been reduced in the field, suggesting that diversity loss may heighten climate change sensitivity. Differences among species within the experimental community in their responses to soil biota and drought suggest that species benefitting from less drought sensitive soil biota may be able to compensate for some of this loss of productivity. Regardless of the plant species and soil origin, further drought eliminated any effects of soil biota on plant growth. Consequently, soil biota may be unable to buffer the effects of drought on primary productivity or other ecosystem functions as extreme events increase in frequency.

Assessing the dual-mycorrhizal status of a widespread tree species as a model for studies on stand biogeochemistry.

Viewing plant species by their mycorrhizal type has explained a range of ecosystem processes. However, mycorrhizal type is confounded with plant phylogeny and the environments in which mycorrhizal partners occur. To circumvent these confounding effects, "dual-mycorrhizal" plant species may be potential models for testing the influence of mycorrhizal type on stand biogeochemistry. To assess their use as models, duality in mycorrhizas within a single host species must be confirmed and factors underlying their variation understood. We surveyed roots, soils, and leaves of mature aspen (Populus tremuloides) across 27 stands in western Canada spanning two biomes: boreal forest and parklands. Aspen roots were mostly ectomycorrhizal with sporadic and rare occurrences of arbuscular mycorrhizas. We further tested whether a climate moisture index predicted abundance of ectomycorrhizal roots (number of ectomycorrhizal root tips m-1 root length) surveyed at two depths (0-20 cm and 20-40 cm) and found that ectomycorrhizal root abundance in subsoils (20-40 cm) was positively related to the index. We subsequently examined the relationships between ectomycorrhizal root abundance, leaf traits, and slow and fast pools of soil organic carbon and nitrogen. The ratio of leaf lignin:N, but not its components, increased along with ectomycorrhizal root abundance in subsoils. Soil carbon and nitrogen pools were independent of ectomycorrhizal root abundance. Our results suggest that (1) categorizing aspen as dual-mycorrhizal may overstate the functional importance of arbuscular mycorrhizas in this species and life stage, (2) water availability influences ectomycorrhizal root abundance, and (3) ectomycorrhizal root abundance coincides with leaf quality.

Resistance of soil biota and plant growth to disturbance increases with plant diversity.

Plant diversity is critical to the functioning of ecosystems, potentially mediated in part by interactions with soil biota. Here, we characterised multiple groups of soil biota across a plant diversity gradient in a long-term experiment. We then subjected soil samples taken along this gradient to drought, freezing and a mechanical disturbance to test how plant diversity affects the responses of soil biota and growth of a focal plant to these disturbances. High plant diversity resulted in soils that were dominated by fungi and associated soil biota, including increased arbuscular mycorrhizal fungi and reduced plant-feeding nematodes. Disturbance effects on the soil biota were reduced when plant diversity was high, resulting in higher growth of the focal plant in all but the frozen soils. These results highlight the importance of plant diversity for soil communities and their resistance to disturbance, with potential feedback effects on plant productivity.

Mechanisms of plant-soil feedback: interactions among biotic and abiotic drivers.

Contents Summary 91 I. Introduction 91 II. Primary PSF mechanisms 91 III. Factors mediating the mechanisms of PSF 93 IV. Conclusions and future directions 94 Acknowledgements 95 Author contributions 95 References 95 SUMMARY: Plant-soil feedback (PSF) occurs when plants alter soil properties that influence the performance of seedlings, with consequent effects on plant populations and communities. Many processes influence PSF, including changes in nutrient availability and the accumulation of natural enemies, mutualists or secondary chemicals. Typically, these mechanisms are investigated in isolation, yet no single mechanism is likely to be completely responsible for PSF as these processes can interact. Further, the outcome depends on which resources are limiting and the other plants and soil biota in the surrounding environment. As such, understanding the mechanisms of PSF and their role within plant communities requires quantification of the interactions among the processes influencing PSF and the associated abiotic and biotic contexts.

Flowering and floral visitation predict changes in community structure provided that mycorrhizas remain intact.

Pollination is critical for plant fitness and population dynamics, yet little attention is paid to the role of flowering and plant-pollinator interactions in structuring plant communities, including community responses to environmental change. Changes in arbuscular mycorrhizal fungi (AMF), nutrient abundances, and plant litter all affect plant access to different resources, and are known regulators of community structure. Each factor can also affect flowering and plant-pollinator interactions, potentially contributing to changes in community structure. To test whether AMF, nutrients, and litter influenced the relationship between pollination and community structure, we conducted a 5-yr field experiment applying fungicide, adding fertilizer, and removing plant litter in native grassland. We measured the distribution of flowers and floral visits among species in year three and linked these measures to changes in plant composition and species richness between years three and five. We hypothesized that an uneven distribution of flowers and visits among species would lead to greater community change, but that the treatments would disrupt this relationship by altering sexual allocation and recruitment. Consistent with our hypothesis, communities with uneven flower distributions exhibited greater changes in community composition and richness under ambient conditions. However, AMF suppression neutralized this relationship and regulated the other treatment effects, highlighting the potential importance of AMF for stabilizing recruitment dynamics. Combined, AMF suppression and nutrient addition caused species losses when few species flowered, likely by compounding stresses for those species. The treatment effects on the relationship between flowering and community composition were more nuanced, but were likely driven by increased competition and altered flowering among species. By contrast, community composition was more stable when visitation rates were uneven among species, irrespective of any treatments. This suggests that some species require high visitation rates to maintain their populations due to greater dependence on sexual reproduction. Combined, these results highlight the importance of flowering and floral visitation to the dynamics of grassland communities. They also suggest that altered recruitment dynamics is a major, yet understudied, mechanism by which environmental change affects communities. Consequently, understanding the effects of environmental change on plant communities will require study of both plant growth and sexual reproduction.

Predicting species establishment using absent species and functional neighborhoods.

Species establishment within a community depends on their interactions with the local environment and resident community. Such environmental and biotic filtering is frequently inferred from functional trait and phylogenetic patterns within communities; these patterns may also predict which additional species can establish. However, differentiating between environmental and biotic filtering can be challenging, which may complicate establishment predictions. Creating a habitat-specific species pool by identifying which absent species within the region can establish in the focal habitat allows us to isolate biotic filtering by modeling dissimilarity between the observed and biotically excluded species able to pass environmental filters. Similarly, modeling the dissimilarity between the habitat-specific species pool and the environmentally excluded species within the region can isolate local environmental filters. Combined, these models identify potentially successful phenotypes and why certain phenotypes were unsuccessful. Here, we present a framework that uses the functional dissimilarity among these groups in logistic models to predict establishment of additional species. This approach can use multivariate trait distances and phylogenetic information, but is most powerful when using individual traits and their interactions. It also requires an appropriate distance-based dissimilarity measure, yet the two most commonly used indices, nearest neighbor (one species) and mean pairwise (all species) distances, may inaccurately predict establishment. By iteratively increasing the number of species used to measure dissimilarity, a functional neighborhood can be chosen that maximizes the detection of underlying trait patterns. We tested this framework using two seed addition experiments in calcareous grasslands. Although the functional neighborhood size that best fits the community's trait structure depended on the type of filtering considered, selecting these functional neighborhood sizes allowed our framework to predict up to 50% of the variation in actual establishment from seed. These results indicate that the proposed framework may be a powerful tool for studying and predicting species establishment.

Plant-soil feedbacks and mycorrhizal type influence temperate forest population dynamics.

Feedback with soil biota is an important determinant of terrestrial plant diversity. However, the factors regulating plant-soil feedback, which varies from positive to negative among plant species, remain uncertain. In a large-scale study involving 55 species and 550 populations of North American trees, the type of mycorrhizal association explained much of the variation in plant-soil feedbacks. In soil collected beneath conspecifics, arbuscular mycorrhizal trees experienced negative feedback, whereas ectomycorrhizal trees displayed positive feedback. Additionally, arbuscular mycorrhizal trees exhibited strong conspecific inhibition at multiple spatial scales, whereas ectomycorrhizal trees exhibited conspecific facilitation locally and less severe conspecific inhibition regionally. These results suggest that mycorrhizal type, through effects on plant-soil feedbacks, could be an important contributor to population regulation and community structure in temperate forests.

Applying the dark diversity concept to nature conservation.

Linking diversity to biological processes is central for developing informed and effective conservation decisions. Unfortunately, observable patterns provide only a proportion of the information necessary for fully understanding the mechanisms and processes acting on a particular population or community. We suggest conservation managers use the often overlooked information relative to species absences and pay particular attention to dark diversity (i.e., a set of species that are absent from a site but that could disperse to and establish there, in other words, the absent portion of a habitat-specific species pool). Together with existing ecological metrics, concepts, and conservation tools, dark diversity can be used to complement and further develop conservation prioritization and management decisions through an understanding of biodiversity relativized by its potential (i.e., its species pool). Furthermore, through a detailed understanding of the population, community, and functional dark diversity, the restoration potential of degraded habitats can be more rigorously assessed and so to the likelihood of successful species invasions. We suggest the application of the dark diversity concept is currently an underappreciated source of information that is valuable for conservation applications ranging from macroscale conservation prioritization to more locally scaled restoration ecology and the management of invasive species.

Species pools, community completeness and invasion: disentangling diversity effects on the establishment of native and alien species.

Invasion should decline with species richness, yet the relationship is inconsistent. Species richness, however, is a product of species pool size and biotic filtering. Invasion may increase with richness if large species pools represent weaker environmental filters. Measuring species pool size and the proportion realised locally (completeness) may clarify diversity-invasion relationships by separating environmental and biotic effects, especially if species' life-history stage and origin are accounted for. To test these relationships, we added seeds and transplants of 15 native and alien species into 29 grasslands. Species pool size and completeness explained more variation in invasion than richness alone. Although results varied between native and alien species, seed establishment and biotic resistance to transplants increased with species pool size, whereas transplant growth and biotic resistance to seeds increased with completeness. Consequently, species pools and completeness represent multiple independent processes affecting invasion; accounting for these processes improves our understanding of invasion.

Macroecology of biodiversity: disentangling local and regional effects.

Contents 404 I. 404 II. 404 III. 405 IV. 406 V. 407 VI. 408 409 References 409 SUMMARY: Macroecology of biodiversity disentangles local and regional drivers of biodiversity by exploring large-scale biodiversity relationships with environmental or biotic gradients, generalizing local biodiversity relationships across regions, or comparing biodiversity patterns among species groups. A macroecological perspective is also important at local scales: a full understanding of local biodiversity drivers, including human impact, demands that regional processes be taken into account. This requires knowledge of which species could inhabit a site (the species pool), including those that are currently absent (dark diversity). Macroecology of biodiversity is currently advancing quickly owing to an unprecedented accumulation of biodiversity data, new sampling techniques and analytical methods, all of which better equip us to face current and future challenges in ecology and biodiversity conservation.

Response to Comment on "Worldwide evidence of a unimodal relationship between productivity and plant species richness".

Tredennick et al. criticize one of our statistical analyses and emphasize the low explanatory power of models relating productivity to diversity. These criticisms do not detract from our key findings, including evidence consistent with the unimodal constraint relationship predicted by the humped-back model and evidence of scale sensitivities in the form and strength of the relationship.

Plant ecology. Worldwide evidence of a unimodal relationship between productivity and plant species richness.

The search for predictions of species diversity across environmental gradients has challenged ecologists for decades. The humped-back model (HBM) suggests that plant diversity peaks at intermediate productivity; at low productivity few species can tolerate the environmental stresses, and at high productivity a few highly competitive species dominate. Over time the HBM has become increasingly controversial, and recent studies claim to have refuted it. Here, by using data from coordinated surveys conducted throughout grasslands worldwide and comprising a wide range of site productivities, we provide evidence in support of the HBM pattern at both global and regional extents. The relationships described here provide a foundation for further research into the local, landscape, and historical factors that maintain biodiversity.

Increased competition does not lead to increased phylogenetic overdispersion in a native grassland.

That competition is stronger among closely related species and leads to phylogenetic overdispersion is a common assumption in community ecology. However, tests of this assumption are rare and field-based experiments lacking. We tested the relationship between competition, the degree of relatedness, and overdispersion among plants experimentally and using a field survey in a native grassland. Relatedness did not affect competition, nor was competition associated with phylogenetic overdispersion. Further, there was only weak evidence for increased overdispersion at spatial scales where plants are likely to compete. These results challenge traditional theory, but are consistent with recent theories regarding the mechanisms of plant competition and its potential effect on phylogenetic structure. We suggest that specific conditions related to the form of competition and trait conservatism must be met for competition to cause phylogenetic overdispersion. Consequently, overdispersion as a result of competition is likely to be rare in natural communities.

Evaluating the relationship between competition and productivity within a native grassland.

Ideas about how plant competition varies with productivity are rooted in classic theories that predict either increasing (Grime) or invariant (Tilman) competition with increasing productivity. Both predictions have received experimental support, although a decade-old meta-analysis supports neither. Attempts to reconcile the conflicting predictions and evidence include: expanding the theory to include other conditions (e.g. stress gradient hypothesis), development of indices to differentiate either the 'intensity' or 'importance' of competition, a focus on resource supply and demand, and explicit recognition that both growth and survival may exhibit different relationships with productivity. To determine which of these theories accurately predict how competition varies with productivity within a native grassland site, we estimated competitive intensity and relative competitive importance using 22 species across the range of productivity naturally occurring within that site. Plant performance was measured as survival and size with and without neighbours and the local environment was quantified according to variability in standing crop, gross water supply, and net water supply. On average, neighbours weakly facilitated seedling survival, but strongly reduced seedling growth. For both seedling survival and growth, relative competitive importance and competitive intensity declined with some measure of productivity; neighbour effects on survival declined with standing crop, while effects on growth declined with gross water supply. These results add to the growing evidence that plant-plant interactions vary among life history components with different life history components contingent upon separate environmental factors. Although the range of productivity measured in this study was not large, our results do not support the theories of Grime or Tilman. However, our results are consistent with the meta-analysis and parts of other theories, although no single theory is capable of explaining the entirety of these results. This suggests that, at least in moderately productive grasslands, new theory needs to be developed.

Context dependence in foraging behaviour of Achillea millefolium.

Context-dependent foraging behaviour is acknowledged and well documented for a diversity of animals and conditions. The contextual determinants of plant foraging behaviour, however, are poorly understood. Plant roots encounter patchy distributions of nutrients and soil fungi. Both of these features affect root form and function, but how they interact to affect foraging behaviour is unknown. We extend the use of the marginal value theorem to make predictions about the foraging behaviour of roots, and test our predictions by manipulating soil resource distribution and inoculation by soil fungi. We measured plant movement as both distance roots travelled and time taken to grow through nutrient patches of varied quality. To do this, we grew Achillea millefolium in the centers of modified pots with a high-nutrient patch and a low-nutrient patch on either side of the plant (heterogeneous) or patch-free conditions (homogeneous). Fungal inoculation, but not resource distribution, altered the time it took roots to reach nutrient patches. When in nutrient patches, root growth decreased relative to homogeneous soils. However, this change in foraging behaviour was not contingent upon patch quality or fungal inoculation. Root system breadth was larger in homogeneous than in heterogeneous soils, until measures were influenced by pot edges. Overall, we find that root foraging behaviour is modified by resource heterogeneity but not fungal inoculation. We find support for predictions of the marginal value theorem that organisms travel faster through low-quality than through high-quality environments, with the caveat that roots respond to nutrient patches per se rather than the quality of those patches.

The Role of Multi Detector Computed Tomographic Imaging Prior to Clinic Consultation in Patients Suspected to Have Lung Cancer.

BACKGROUND: Scanning patients early in their diagnostic journey helps in differentiating benign from malignant aetiology. There is increasing pressure on diagnostic practices for rapid diagnoses and thereby early commencement of treatment in patients suspected to have lung cancer (LC). In our practice, multi detector computed tomography (MDCT) imaging is performed in selected patients referred to the LC service prior to them seeing a chest physician in the LC clinic. This study evaluates the role of such practice and reviews its potential impact on LC services. METHODS: Prospective review of our practice from January 2007 to Apr 2007 was performed. Consecutive patients referred to the service with suspected LC were included. Chest radiograph (CXR) report and clinical information from general practitioners were reviewed and graded as high, medium or low risk for presence of LC. Patients with sufficient clinical and/or radiological concern underwent MDCT imaging prior to their clinic. Combined risk scores and modified risk scores were formulated and assessed against MDCT findings. RESULTS: A total of 139 patients were referred to the service, 124 of these had pre-clinic MDCT. Fifty-three patients (43%) had malignancy, 39 (31%) had non-malignant significant abnormalities, 17 (14%) had other incidental findings and 15 (12%) were normal. Modified combined risk score was the best predictor of presence of cancer. CONCLUSION: Pre-clinic MDCT scanning in patients with suspected LC is feasible and has a promising role in the modern care of LC patients. It also empowers physicians with additional information at the primary consultation.

Thoracic ultrasound prior to medical thoracoscopy improves pleural access and predicts fibrous septation.

BACKGROUND AND OBJECTIVE: Medical thoracoscopy (MT) is indicated for the investigation of unexplained pleural exudates. Not all MT units create artificial pneumothoraces because of time. Difficult pleural space access and thick fibrous adhesions may prevent MT and pleurodesis, respectively. The potential role of thoracic ultrasound (TUS) pre-MT has not been fully evaluated. We hypothesized TUS would reduce failure to access the pleural space and enable detection of thick fibrous adhesions. METHODS: Thirty patients underwent single port MT consecutively for investigation of pleural exudates without pre-MT TUS over a 6-month period. Over the following 6 months, 30 consecutive patients underwent TUS immediately prior to MT. Pleural access rate and thick fibrous adhesion detection at both MT and TUS were recorded. RESULTS: In the non-TUS cohort, pleural space access failure occurred in 16.7% (leading to five extra procedures), versus no failures in the TUS cohort (P = 0.0522). There were no differences in prevalence of MT fibrous adhesions between cohorts. TUS identified all cases of fibrous septation versus only 12.5% identified by CT in the non-TUS cohort (P = 0.001). All identified cases of thick fibrous septation on TUS did not receive pleurodesis at MT. TUS detected useful ancillary features in 43% of cases. CONCLUSIONS: A strong trend to reduction in single port MT pleural access failure was noted with pre-MT TUS thus reducing extra procedures and the need for artificial pneumothoraces. Pre-MT TUS also reliably detects thick fibrous adhesions at MT. TUS may also detect useful ancillary features. This study provides a rationale for ultrasound-guided single port MT if a pneumothorax is not created.

Current status of medical pleuroscopy.

Medical pleuroscopy (MP) offers a safe and minimally invasive tool for interventional pulmonologists. It allows diagnosis of unexplained effusion, while at the same time allowing drainage and pleurodesis. It can also help in the diagnosis of diffuse interstitial disease or associated peripheral lung abnormality in the presence of effusion. It can have a therapeutic role in pneumothorax and hyperhidrosis or chronic pancreatic pain. This article reviews the technical aspects and range of applications of MP.