Precipitation patterns including the magnitude, timing, and seasonality of rainfall are predicted to undergo substantial alterations in arid regions in the future, and desert organisms may be more responsive to such changes than to shifts in only mean annual rainfall. Soil biocrust communities (consisting of cyanobacteria, lichen, and mosses) are ubiquitous to desert ecosystems, play an array of ecological roles, and display a strong sensitivity to environmental changes. Crust mosses are particularly responsive to changes in precipitation and exhibit rapid declines in biomass and mortality following the addition of small rainfall events. Further, loss of the moss component in biocrusts leads to declines in crust structure and function. In this study, we sought to understand the physiological responses of the widespread and often dominant biocrust moss Syntrichia caninervis to alterations in rainfall. Moss samples were collected during all four seasons and exposed to two rainfall event sizes and three desiccation period (DP) lengths. A carbon balance approach based on single precipitation events was used to define the carbon gain or loss during a particular hydration period. Rainfall event size was the strongest predictor of carbon balance, and the largest carbon gains were associated with the largest precipitation events. In contrast, small precipitation events resulted in carbon deficits for S. caninervis. Increasing the length of the DP prior to an event resulted in reductions in carbon balance, probably because of the increased energetic cost of hydration following more intense bouts of desiccation. The season of collection (i.e., physiological status of the moss) modulated these responses, and the effects of DP and rainfall on carbon balance were different in magnitude (and often in sign) for different seasons. In particular, S. caninervis displayed higher carbon balances in the winter than in the summer, even for events of identical size. Overall, our results suggest that annual carbon balance and survivorship in biocrust mosses are largely driven by precipitation, and because of the role mosses play in biocrusts, changes in intra-annual precipitation patterns can have implications for hydrology, soil stability, and nutrient cycling in dryland systems.
With global climate change, water scarcity threatens whole agro/ecosystems. The desert moss Syntrichia caninervis, an extremophile, offers novel insights into surviving desiccation and heat. The sequenced S. caninervis genome consists of 13 chromosomes containing 16 545 protein-coding genes and 2666 unplaced scaffolds. Syntenic relationships within the S. caninervis and Physcomitrella patens genomes indicate the S. caninervis genome has undergone a single whole genome duplication event (compared to two for P. patens) and evidence suggests chromosomal or segmental losses in the evolutionary history of S. caninervis. The genome contains a large sex chromosome composed primarily of repetitive sequences with a large number of Copia and Gypsy elements. Orthogroup analyses revealed an expansion of ELIP genes encoding proteins important in photoprotection. The transcriptomic response to desiccation identified four structural clusters of novel genes. The genomic resources established for this extremophile offer new perspectives for understanding the evolution of desiccation tolerance in plants.
PREMISE Desiccation tolerance (DT) is a widespread phenomenon among land plants, and variable ecological strategies for DT are likely to exist. Using Syntrichia caninervis, a dryland moss and model system used in DT studies, we hypothesized that DT is lowest in juvenile (protonemal) tissues, highest in asexual reproductive propagules (gemmae), and intermediate in adults (shoots). We tested the long‐standing hypothesis of an inherent constitutive strategy of DT in this species. METHODS Plants were rapidly dried to levels of equilibrating relative humidity (RHeq) ranging from 0 to 93%. Postrehydration recovery was assessed using chlorophyll fluorescence, regeneration rates, and visual tissue damage. For each life phase, we estimated the minimum rate of drying (RoDmin) at RHeq = 42% that did not elicit damage 24 h postrehydration. RESULTS DT strategy varied with life phase, with adult shoots having the lowest RoDmin (10‒25 min), followed by gemmae (3‒10 h) and protonema (14‒20 h). Adult shoots exhibited no detectable damage 24 h postrehydration following a rapid‐dry only at the highest RHeq used (93%), but when dried to lower RHs the response declined to <50% of control fluorescence values. Notably, immediately following rehydration (0 h postrehydration), shoots were damaged below control levels of fluorescence regardless of the RHeq, thus implicating damage. CONCLUSIONS Life phases of the moss S. caninervis had a range of strategies from near constitutive (adult shoots) to demonstrably inducible (protonema). A new response variable for assessing degree of DT is introduced as the minimum rate of drying from which full recovery occurs.
Fluctuations in mean annual precipitation (MAP) will strongly influence the ecology of dryland ecosystems in the future, yet, because individual precipitation events drive growth and resource availability for many dryland organisms, changes in intra-annual precipitation may disproportionately influence future dryland processes. This work examines the hypothesis that intra-annual precipitation changes will drive dryland productivity to a greater extent than changes to MAP. To test this hypothesis, we created a physiology-based model to predict the effects of precipitation change on a widespread biocrust moss that regulates soil structure, water retention, and nutrient cycling in drylands. First, we used the model to examine moss productivity over the next 100 years driven by alterations in MAP by ± 10, 20 and 30%, and changes in intra-annual precipitation (event size and frequency). Productivity increased as a function of MAP, but differed among simulations where intra-annual precipitation was manipulated under constant MAP. Supporting our hypothesis, this demonstrates that, even if MAP does not change, changes in the features of individual precipitation events can strongly influence long-term performance. Second, we used the model to examine 100-year productivity based on projected dryland precipitation from published global and regional models. These simulations predicted 25-63% reductions in productivity and increased moss mortality rates, declines that will likely alter water and nutrient cycling in dryland ecosystems. Intra-annual precipitation in model-based simulations was a stronger predictor of productivity compared to MAP, further supporting our hypothesis, and illustrating that intra-annual precipitation patterns may dominate dryland responses to altered precipitation in a future climate.
Since the late Cambrian era, bryophytes have been shaping terrestrial ecosystems through unique and diverse suites of anatomical, physiological, and morphological traits. In this review we highlight historical and recent work in bryophyte functional ecology, with an emphasis on knowledge gaps and opportunities for future work. While we cannot always avoid the temptation to contrast with tracheophyte (especially angiosperm) studies, our aim is to de-center that perspective in favor of a more universal understanding of functional land plant ecology. We therefore center our description on three core aspects of bryophytes that are poorly represented in tracheophyte studies: (I) dynamic water content (including poikilohydry and desiccation tolerance), (II) multiple scales of interaction with environment, and (III) reproduction and life history. We also highlight the diverse and wide-ranging influence bryophytes have on ecosystem processes, including primary productivity, nutrient cycling, hydrology, and ecological interactions with other species. Furthermore, while the study of bryophyte functional traits has rapidly grown in the past decade, important gaps in phylogenetic and geographic coverage persist and constrain the development of a more universal land plant functional ecology theory.
Plant functional trait analyses have focused almost exclusively on vascular plants, but bryophytes comprise ancient and diverse plant lineages that have widespread global distributions and important ecological functions in terrestrial ecosystems. We examined a diverse clade of dryland mosses, Syntrichia, and studied carbon balance during a precipitation event (C‐balance), a functional trait related to physiological functioning, desiccation tolerance, survival, and ecosystem carbon and nitrogen cycling. We examined variability in C‐balance among 14 genotypes of Syntrichia and measured an additional 10 physiological and 13 morphological traits at the cell, leaf, shoot, and clump level. C‐balance varied 20‐fold among genotypes, and highest C‐balances were associated with long, narrow leaves with awns, and small cells with thick cell walls, traits that may influence water uptake and retention during a precipitation event. Ordination analyses revealed that the axis most strongly correlated with C‐balance included the maximum chlorophyll fluorescence, Fm, indicating the importance of photosystem II health for C exchange. C‐balance represents a key functional trait in bryophytes, but its measurement is time intensive and not feasible to measure on large scales. We propose two models (using physiological and morphological traits) to predict C‐balance, whereby identifying simpler to measure traits for trait databases.
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