Symbionts out of sync: Decoupled physiological responses are widespread and ecologically important in lichen associations.

A core vulnerability in symbioses is the need for coordination between the symbiotic partners, which are often assumed to be closely physiologically integrated. We critically re-examine this assumed integration between symbionts in lichen symbioses, recovering a long overlooked yet fundamental physiological asymmetry in carbon balance. We examine the physiological, ecological, and transcriptional basis of this asymmetry in the lichen Evernia mesomorpha. This carbon balance asymmetry depends on hydration source and aligns with climatic range limits. Differences in gene expression across the E. mesomorpha symbiosis suggest that the physiologies of the primary lichen symbionts are decoupled. Furthermore, we use gas exchange data to show that asymmetries in carbon balance are widespread and common across evolutionarily disparate lichen associations. Using carbon balance asymmetry as an example, we provide evidence for the wide-ranging importance of physiological asymmetries in symbioses.

Carbon-concentrating mechanisms are a key trait in lichen ecology and distribution.

Carbon-concentrating mechanisms (CCMs) are a widespread phenomenon in photosynthetic organisms. In vascular plants, the evolution of CCMs ([C44-carbon compound] and crassulacean acid metabolism [CAM]) is associated with significant shifts, most often to hot, dry and bright, or aquatic environments. If and how CCMs drive distributions of other terrestrial photosynthetic organisms, remains little studied. Lichens are ecologically important obligate symbioses between fungi and photosynthetic organisms. The primary photosynthetic partner in these symbioses can include CCM-presenting cyanobacteria (as carboxysomes), CCM-presenting green algae (as pyrenoids) or green algae lacking any CCM. We use an extensive dataset of lichen communities from eastern North America, spanning a wide climatic range, to test the importance of CCMs as predictors of lichen ecology and distribution. We show that the presence or absence of CCMs leads to opposite responses to temperature and precipitation in green algal lichens, and different responses in cyanobacterial lichens. These responses contrast with our understanding of lichen physiology, whereby CCMs mitigate carbon limitation by water saturation at the cost of efficient use of vapor hydration. This study demonstrates that CCM status is a key functional trait in obligate lichen symbioses, equivalent in importance to its role in vascular plants, and central for studying present and future climate responses.

Lichen ecophysiology in a changing climate.

Lichens are one of the most iconic and ubiquitous symbioses known, widely valued as indicators of environmental quality and, more recently, climate change. Our understanding of lichen responses to climate has greatly expanded in recent decades, but some biases and constraints have shaped our present knowledge. In this review we focus on lichen ecophysiology as a key to predicting responses to present and future climates, highlighting recent advances and remaining challenges. Lichen ecophysiology is best understood through complementary whole-thallus and within-thallus scales. Water content and form (vapor or liquid) are central to whole-thallus perspectives, making vapor pressure differential (VPD) a particularly informative environmental driver. Responses to water content are further modulated by photobiont physiology and whole-thallus phenotype, providing clear links to a functional trait framework. However, this thallus-level perspective is incomplete without also considering within-thallus dynamics, such as changing proportions or even identities of symbionts in response to climate, nutrients, and other stressors. These changes provide pathways for acclimation, but their understanding is currently limited by large gaps in our understanding of carbon allocation and symbiont turnover in lichens. Lastly, the study of lichen physiology has mainly prioritized larger lichens at high latitudes, producing valuable insights but underrepresenting the range of lichenized lineages and ecologies. Key areas for future work include improving geographic and phylogenetic coverage, greater emphasis on VPD as a climatic factor, advances in the study of carbon allocation and symbiont turnover, and the incorporation of physiological theory and functional traits in our predictive models.

Evolutionary biology of lichen symbioses.

Lichens are the symbiotic outcomes of open, interspecies relationships, central to which are a fungus and a phototroph, typically an alga and/or cyanobacterium. The evolutionary processes that led to the global success of lichens are poorly understood. In this review, we explore the goods and services exchange between fungus and phototroph and how this propelled the success of both symbiont and symbiosis. Lichen fungal symbionts count among the only filamentous fungi that expose most of their mycelium to an aerial environment. Phototrophs export carbohydrates to the fungus, which converts them to specific polyols. Experimental evidence suggests that polyols are not only growth and respiratory substrates but also play a role in anhydrobiosis, the capacity to survive desiccation. We propose that this dual functionality is pivotal to the evolution of fungal symbionts, enabling persistence in environments otherwise hostile to fungi while simultaneously imposing costs on growth. Phototrophs, in turn, benefit from fungal protection from herbivory and light stress, while appearing to exert leverage over fungal sex and morphogenesis. Combined with the recently recognized habit of symbionts to occur in multiple symbioses, this creates the conditions for a multiplayer marketplace of rewards and penalties that could drive symbiont selection and lichen diversification.

Differential retention contributes to racial/ethnic disparity in U.S. academia.

Several racial and ethnic identities are widely understood to be under-represented within academia, however, actual quantification of this under-representation is surprisingly limited. Challenges include data availability, demographic inertia and identifying comparison points. We use de-aggregated data from the U.S. National Science Foundation to construct a null model of ethnic and racial representation in one of the world's largest academic communities. Making comparisons between our model and actual representation in academia allows us to measure the effects of retention (while controlling for recruitment) at different academic stages. We find that, regardless of recruitment, failed retention contributes to mis-representation across academia and that the stages responsible for the largest disparities differ by race and ethnicity: for Black and Hispanic scholars this occurs at the transition from graduate student to postdoctoral researcher whereas for Native American/Alaskan Native and Native Hawaiian/Pacific Islander scholars this occurs at transitions to and within faculty stages. Even for Asian and Asian-Americans, often perceived as well represented, circumstances are complex and depend on choice of baseline. Our findings demonstrate that while recruitment continues to be important, retention is also a pervasive barrier to proportional representation. Therefore, strategies to reduce mis-representation in academia must address retention. Although our model does not directly suggest specific strategies, our framework could be used to project how representation in academia might change in the long-term under different scenarios.

Let's move out together: a framework for the intersections between movement and mutualism.

Movement is a widespread behavior across organisms and is driven in part by interspecific interactions. Generally, negative interspecific interactions (such as competition and natural enemies) are more often studied in the context of movement than positive interactions (mutualism). Mutualistic relationships are incredibly common, yet only a subset are studied in the context of movement (transportation mutualisms). Overall, the costs and benefits that an individual experiences are shaped both by their movement behavior and their mutualistic relationships, as well as the intersection between these. Here we argue that the intersection between movement behavior and mutualistic relationships is understudied, and we present a conceptual framework to synthesize the links between movement and mutualisms and give examples of species that exhibit each. Our framework serves both to highlight the ways that mutualism can shape movement (and vice versa) and to draw parallels across different organisms (enabling a more abstract perspective of these biological systems, complementing the system-focused perspective). Finally, we show how considering movement in light of mutualisms (and vice versa) presents a number of new research questions to be answered by each empirical and theoretical approach going forward.

Soil invertebrate diversity loss and functional changes in temperate forest soils replaced by exotic pine plantations.

The global expansion of tree plantations is often claimed to have positive effects for mitigating global warming, preventing soil erosion, and reducing biodiversity loss. However, questions remain unanswered about the impacts of plantations on belowground diversity and soil properties. Here, we examine how forestry plantations of exotic trees affect critical soil functions and the composition of invertebrate assemblages, by comparing invertebrate diversity and soil physico-chemical properties between non-native Pinus radiata plantations, and nearby native forests in a region of extensive plantation activity in south-central Chile. We quantified differences in diversity, abundance, and community composition of soil invertebrates, as well as fundamental soil properties such as soil water content, water infiltration, nutrient status, and pH. We show that in this landscape mosaic of native forest and plantations, both soil invertebrate communities and physical soil properties differed significantly between systems, despite similar soil origins and topographies. We found a significant loss of soil carbon and a major reduction in taxonomic and functional diversity of soil invertebrates in pine plantation sites. Soil biotic and abiotic characteristics of plantations differed significantly from native forests in plantation-dominated south-central Chile, with profound consequences for ecosystem processes and resilience to future climate change.

Rapid nitrogen fixation by canopy microbiome in tropical forest determined by both phosphorus and molybdenum.

Biological nitrogen fixation is critical for the nitrogen cycle of tropical forests, yet we know little about the factors that control the microbial nitrogen fixers that colonize the microbiome of leaves and branches that make up a forest canopy. Forest canopies are especially prone to nutrient limitation because they are (1) disconnected from soil nutrient pools and (2) often subject to leaching. Earlier studies have suggested a role of phosphorus and molybdenum in controlling biological N-fixation rates, but experimental confirmation has hitherto been unavailable. Here we present the results of a manipulation of canopy nutrient availability. Our findings demonstrate a primary role of phosphorus in constraining overall N fixation by canopy cyanobacteria, but also a secondary role of molybdenum in determining per-cell fixation rates. A conservative evaluation suggests that canopy fixation can contribute to significant N fluxes at the ecosystem level, especially as bursts following atmospheric inputs of nutrient-rich dust.

Vertically challenged: How disease suppresses Daphnia vertical migration behavior.

Parasitic infections are increasingly recognized as influential forces in the migratory behaviors of hosts ranging from butterflies to whales. In aquatic zooplankton, diel vertical migrations (DVMs) are among the most recurrent behaviors with implications for predator-prey interactions, nutrient cycling, and energy flow, yet how parasitism affects such migrations remains an open question. Here, we tested the effects of sporangia cluster disease (SCD) on DVM of the large-bodied Daphnia pulicaria, which is often considered a key component of lake food webs. By collecting depth-specific zooplankton samples across diel cycles, between years, and among lakes, we show that infection is associated with strong inhibition of host DVM; while all Daphnia tended to occur deeper during the day, uninfected Daphnia and especially gravid individuals migrated to shallower waters at night. In contrast, infected hosts-which could comprise 40% of the population-were more likely to remain deep regardless of time of day. Among infected hosts, the intensity of SCD (sporangia count per host) predicted the degree of DVM inhibition. These observations-coupled with lab-based assays showing that infected hosts exhibited fewer swimming movements and persisted at lower depths than uninfected conspecifics-suggest that parasite-induced inhibition of DVM is a "sickness behavior" resulting from increasing morbidity and energy depletion as the infection intensifies toward host death. Considering the importance of large-bodied Daphnia as regulators of water clarity and prey for fishes, parasite-induced alterations of host migratory behavior have broad potential to affect the redistribution of energy and nutrients within lake ecosystems.

Morphogeometric Approaches to Non-vascular Plants.

Morphometric analysis of organisms has undergone a dramatic renaissance in recent years, embracing a range of novel computational and imaging techniques to provide new approaches to phenotypic characterization. These innovations have often developed piece-meal, and may reflect the taxonomic specializations and biases of their creators. In this review, we aim to provide a brief introduction to applications and applicability of modern morphometrics to non-vascular land plants, an often overlooked but evolutionarily and ecologically important group. The scale and physiology of bryophytes (mosses, liverworts, and hornworts) differ in important and informative ways from more "traditional" model plants, and their inclusion has the potential to powerfully broaden perspectives in plant morphology. In particular we highlight three areas where the "bryophytic perspective" shows considerable inter-disciplinary potential: (i) bryophytes as models for intra-specific and inter-specific phenotypic variation, (ii) bryophyte growth-forms as areas for innovation in architectural modularity, and (iii) bryophytes as models of ecophysiological integration between organs, individuals, and stands. We suggest that advances should come from two-way dialog: the translation and adoption of techniques recently developed for vascular plants (and other organisms) to bryophytes and the use of bryophytes as model systems for the innovation of new techniques and paradigms in morphogeometric approaches.

Easy Come, Easy Go: Capillary Forces Enable Rapid Refilling of Embolized Primary Xylem Vessels.

Protoxylem plays an important role in the hydraulic function of vascular systems of both herbaceous and woody plants, but relatively little is known about the processes underlying the maintenance of protoxylem function in long-lived tissues. In this study, embolism repair was investigated in relation to xylem structure in two cushion plant species, Azorella macquariensis and Colobanthus muscoides, in which vascular water transport depends on protoxylem. Their protoxylem vessels consisted of a primary wall with helical thickenings that effectively formed a pit channel, with the primary wall being the pit channel membrane. Stem protoxylem was organized such that the pit channel membranes connected vessels with paratracheal parenchyma or other protoxylem vessels and were not exposed directly to air spaces. Embolism was experimentally induced in excised vascular tissue and detached shoots by exposing them briefly to air. When water was resupplied, embolized vessels refilled within tens of seconds (excised tissue) to a few minutes (detached shoots) with water sourced from either adjacent parenchyma or water-filled vessels. Refilling occurred in two phases: (1) water refilled xylem pit channels, simplifying bubble shape to a rod with two menisci; and (2) the bubble contracted as the resorption front advanced, dissolving air along the way. Physical properties of the protoxylem vessels (namely pit channel membrane porosity, hydrophilic walls, vessel dimensions, and helical thickenings) promoted rapid refilling of embolized conduits independent of root pressure. These results have implications for the maintenance of vascular function in both herbaceous and woody species, because protoxylem plays a major role in the hydraulic systems of leaves, elongating stems, and roots.

Growth responses of the mangrove Avicennia marina to salinity: development and function of shoot hydraulic systems require saline conditions.

BACKGROUND AND AIMS: Halophytic eudicots are characterized by enhanced growth under saline conditions. This study combines physiological and anatomical analyses to identify processes underlying growth responses of the mangrove Avicennia marina to salinities ranging from fresh- to seawater conditions. METHODS: Following pre-exhaustion of cotyledonary reserves under optimal conditions (i.e. 50% seawater), seedlings of A. marina were grown hydroponically in dilutions of seawater amended with nutrients. Whole-plant growth characteristics were analysed in relation to dry mass accumulation and its allocation to different plant parts. Gas exchange characteristics and stable carbon isotopic composition of leaves were measured to evaluate water use in relation to carbon gain. Stem and leaf hydraulic anatomy were measured in relation to plant water use and growth. KEY RESULTS: Avicennia marina seedlings failed to grow in 0-5% seawater, whereas maximal growth occurred in 50-75% seawater. Relative growth rates were affected by changes in leaf area ratio (LAR) and net assimilation rate (NAR) along the salinity gradient, with NAR generally being more important. Gas exchange characteristics followed the same trends as plant growth, with assimilation rates and stomatal conductance being greatest in leaves grown in 50-75% seawater. However, water use efficiency was maintained nearly constant across all salinities, consistent with carbon isotopic signatures. Anatomical studies revealed variation in rates of development and composition of hydraulic tissues that were consistent with salinity-dependent patterns in water use and growth, including a structural explanation for low stomatal conductance and growth under low salinity. CONCLUSIONS: The results identified stem and leaf transport systems as central to understanding the integrated growth responses to variation in salinity from fresh- to seawater conditions. Avicennia marina was revealed as an obligate halophyte, requiring saline conditions for development of the transport systems needed to sustain water use and carbon gain.

Ecosystem properties self-organize in response to a directional fog-vegetation interaction.

Feedbacks between vegetation and resource inputs can lead to the local, self-organization of ecosystem properties. In particular, feedbacks in response to directional resources (e.g., coastal fog, slope runoff) can create complex spatial patterns, such as vegetation banding. Although similar feedbacks are thought to be involved in the development of ecosystems, clear empirical examples are rare. We created a simple model of a fog-influenced, temperate rainforest in central Chile, which allows the comparison of natural banding patterns to simulations of various putative mechanisms. We show that only feedbacks between plants and fog were able to replicate the characteristic distributions of vegetation, soil water, and soil nutrients observed in field transects. Other processes, such as rainfall, were unable to match these diagnostic distributions. Furthermore, fog interception by windward trees leads to increased downwind mortality, leading to progressive extinction of the leeward edge. This pattern of ecosystem development and decay through self-organized processes illustrates, on a relatively small spatial and temporal scale, the patterns predicted for ecosystem evolution.

Water redistribution determines photosynthetic responses to warming and drying in two polar mosses.

Predicting impacts of climate change requires an understanding of the sensitivity of species to temperature, including conflated changes in humidity. Physiological responses to temperature and clump-to-air vapour pressure difference (VPD) were compared in two Antarctic moss species, Ceratodon purpureus (Hedw.) Brid. and Schistidium antarctici (Cardot) L.I. Savicz & Smirnova. Temperatures from 8 to 24°C had no significant effects on photosynthesis or recovery from drying, whereas high VPD accelerated drying. In Schistidium, which lacks internal conduction structures, shoots dried more slowly than the clump, and photosynthesis ceased at high shoot relative water content (RWC), behaviour consistent with a strategy of drought avoidance although desiccation tolerant. In contrast, shoots of Ceratodon have a central vascular core, but dried more rapidly than the clump. These results imply that cavitation of the hydroid strand enables hydraulic isolation of extremities during rapid drying, effectively slowing water loss from the clump. Ceratodon maintained photosynthetic activity during drying to lower shoot RWC than Schistidium, consistent with a strategy of drought tolerance. These ecophysiological characteristics may provide a functional explanation for the differential distribution of Schistidium and Ceratodon along moisture gradients in Antarctica. Thus, predicting responses of non-vascular vegetation to climate change at high latitudes requires greater focus on VPD and hydraulics than temperature.

Leaks in the pipeline: separating demographic inertia from ongoing gender differences in academia.

Identification of the causes underlying the under-representation of women and minorities in academia is a source of ongoing concern and controversy. This is a critical issue in ensuring the openness and diversity of academia; yet differences in personal experiences and interpretations have mired it in controversy. We construct a simple model of the academic career that can be used to identify general trends, and separate the demographic effects of historical differences from ongoing biological or cultural gender differences. We apply the model to data on academics collected by the National Science Foundation (USA) over the past three decades, across all of science and engineering, and within six disciplines (agricultural and biological sciences, engineering, mathematics and computer sciences, physical sciences, psychology, and social sciences). We show that the hiring and retention of women in academia have been affected by both demographic inertia and gender differences, but that the relative influence of gender differences appears to be dwindling for most disciplines and career transitions. Our model enables us to identify the two key non-structural bottlenecks restricting female participation in academia: choice of undergraduate major and application to faculty positions. These transitions are those in greatest need of detailed study and policy development.

Dining on disease: how interactions between infection and environment affect predation risk.

Despite growing interest in ecological interactions between predators and pathogens, few studies have experimentally examined the consequences of infection for host predation risk or how environmental conditions affect this relationship. Here we combined mesocosm experiments, in situ foraging data, and broad-scale lake surveys to evaluate (1) the effects of chytrid infection (Polycaryum laeve) on susceptibility of Daphnia to fish predators and (2) how environmental characteristics moderate the strength of this interaction. In mesocosms, bluegill preferred infected Daphnia 2-5 times over uninfected individuals. Among infected Daphnia, infection intensity was a positive predictor of predation risk, whereas carapace size and fecundity increased predation on uninfected individuals. Wild-caught yellow perch and bluegill from in situ foraging trials exhibited strong selectivity for infected Daphnia (3-10 times over uninfected individuals). In mesocosms containing water high in dissolved organic carbon (DOC), however, selective predation on infected Daphnia was eliminated. Correspondingly, lakes that supported chytrid infections had higher DOC levels and lower light penetration. Our results emphasize the strength of interactions between parasitism and predation while highlighting the moderating influence of water color. P. laeve increases the conspicuousness and predation risk of Daphnia; as a result, infected Daphnia occur predominantly in environments with characteristics that conceal their elevated visibility.