Contrasting shrub and grass hydraulic responses to experimental drought.

Whole-plant hydraulics provide important information about responses to water limitation and can be used to understand how plant communities may change in a drier climate when measured on multiple species. Here, we measured above- and belowground hydraulic traits in Cornus drummondii, an encroaching shrub within North American tallgrass prairies, and Andropogon gerardii, a dominant C4 grass, to assess the potential hydraulic responses to future drought as this region undergoes woody expansion. Shelters that reduced precipitation by 50% and 0% were built over shrubs and grasses growing in sites that are burned at 1-year and 4-year frequencies. We then measured aboveground (Kshoot), belowground (Kroot), and whole-plant maximum hydraulic conductance (Kplant) in C. drummondii and Kroot in A. gerardii. We also measured vulnerability to embolism (P50) in C. drummondii stems. Overall, we show that: (1) A. gerardii had substantially greater Kroot than C. drummondii; (2) belowground hydraulic functioning was linked with aboveground processes; (3) above- and belowground C. drummondii hydraulics were not negatively impacted by the rainfall reductions imposed here. These results suggest that a multi-year drought will not ameliorate rates of woody expansion and highlight key differences in aboveground and belowground hydraulics for dominant species within the same ecosystem.

Linking stem rehydration kinetics to hydraulic traits using a novel method and mechanistic model.

BACKGROUND: Despite the recognized importance of hydraulic capacitance as a mechanism used by plants to maintain hydraulic functioning during high transpiration, characterizing the dynamics of capacitance remains a challenge. METHODS: We used a novel 'two-balance method' to investigate relationships between stem rehydration kinetics and other hydraulic traits in multiple tree species, and we developed a model to explore stem rehydration kinetics further. KEY RESULTS: We found that: (1) rehydration time constants and the amount of water uptake occurring during rehydration differed significantly across species; (2) time constants did not change with declining water potential (Ψ), while water uptake increased at lower Ψ in some species; (3) longer time constants were associated with lower wood density, higher capacitance and less negative stem pressures causing 50 % loss of hydraulic conductivity (P50); (4) greater water uptake occurred in stems with lower wood density and less negative P50 values; and (5) the model could estimate the total hydraulic resistance of the rehydration path, which cannot be measured directly. CONCLUSIONS: Overall, the two-balance method can be used to examine rehydration dynamics quickly and thoroughly in detached woody stems. This method has the potential to improve our understanding of how capacitance functions across tree species, which is an often-overlooked component of whole-plant hydraulics.

At least it is a dry cold: the global distribution of freeze-thaw and drought stress and the traits that may impart poly-tolerance in conifers.

Conifers inhabit some of the most challenging landscapes where multiple abiotic stressors (e.g., aridity, freezing temperatures) often co-occur. Physiological tolerance to multiple stressors ('poly-tolerance') is thought to be rare because exposure to one stress generally limits responses to another through functional trade-offs. However, the capacity to exhibit poly-tolerance may be greater when combined abiotic stressors have similar physiological impacts, such as the disruption of hydraulic function imposed by drought or freezing. Here, we reviewed empirical data in light of theoretical expectations for conifer adaptations to drought and freeze-thaw cycles with particular attention to hydraulic traits of the stem and leaf. Additionally, we examined the commonality and spatial distribution of poly-stress along indices of these combined stressors. We found that locations with the highest values of our poly-stress index (PSi) are characterized by moderate drought and moderate freeze-thaw, and most of the global conifer distribution occupies areas of moderate poly-stress. Among traits examined, we found diverse responses to the stressors. Turgor loss point did not correlate with freeze-thaw or drought stress individually, but did with the PSi, albeit inverse to what was hypothesized. Leaf mass per area was more strongly linked with drought stress than the poly-stress and not at all with freeze-thaw stress. In stems, the water potential causing 50% loss of hydraulic conductivity became more negative with increasing drought stress and poly-stress but did not correlate with freeze-thaw stress. For these traits, we identified a striking lack of coverage for substantial portions of species ranges, particularly at the upper boundaries of their respective PSis, demonstrating a critical gap in our understanding of trait prevalence and plasticity along these stress gradients. Future research should investigate traits that confer tolerance to both freeze-thaw and drought stress in a wide range of species across broad geographic scales.

N and P constrain C in ecosystems under climate change: Role of nutrient redistribution, accumulation, and stoichiometry.

We use the Multiple Element Limitation (MEL) model to examine responses of 12 ecosystems to elevated carbon dioxide (CO2 ), warming, and 20% decreases or increases in precipitation. Ecosystems respond synergistically to elevated CO2 , warming, and decreased precipitation combined because higher water-use efficiency with elevated CO2 and higher fertility with warming compensate for responses to drought. Response to elevated CO2 , warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO2 and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C-nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO2 and climate change.

Spatio-temporal differences in leaf physiology are associated with fire, not drought, in a clonally integrated shrub.

In highly disturbed environments, clonality facilitates plant survival via resprouting after disturbance, resource sharing among interconnected stems and vegetative reproduction. These traits likely contribute to the encroachment of deep-rooted clonal shrubs in tallgrass prairie. Clonal shrubs have access to deep soil water and are typically thought of as relatively insensitive to environmental variability. However, how leaf physiological traits differ among stems within individual clonal shrubs (hereafter 'intra-clonal') in response to extreme environmental variation (i.e. drought or fire) is unclear. Accounting for intra-clonal differences among stems in response to disturbance is needed to more accurately parameterize models that predict the effects of shrub encroachment on ecosystem processes. We assessed intra-clonal leaf-level physiology of the most dominant encroaching shrub in Kansas tallgrass prairie, Cornus drummondii, in response to precipitation and fire. We compared leaf gas exchange rates from the periphery to centre within shrub clones during a wet (2015) and extremely dry (2018) year. We also compared leaf physiology between recently burned shrubs (resprouts) with unburned shrubs in 2018. Resprouts had higher gas exchange rates and leaf nitrogen content than unburned shrubs, suggesting increased rates of carbon gain can contribute to recovery after fire. In areas recently burned, resprouts had higher gas exchange rates in the centre of the shrub than the periphery. In unburned areas, leaf physiology remained constant across the growing season within clonal shrubs (2015 and 2018). Results suggest single measurements within a shrub are likely sufficient to parameterize models to understand the effects of shrub encroachment on ecosystem carbon and water cycles, but model parameterization may require additional complexity in the context of fire.

An assessment of diurnal water uptake in a mesic prairie: evidence for hydraulic lift?

Hydraulic lift, the passive movement of water through plant roots from wet to dry soil, is an important ecohydrological process in a wide range of water-limited ecosystems. This phenomenon may also alter plant functioning, growth, and survival in mesic grasslands, where soil moisture is spatially and temporally variable. Here, we monitored diurnal changes in the isotopic signature of soil and plant xylem water to assess (1) whether hydraulic lift occurs in woody and herbaceous tallgrass prairie species (Rhus glabra, Amorpha canescens, Vernonia baldwinii, and Andropogon gerardii), (2) if nocturnal transpiration or grazing by large ungulates limits hydraulic lift, and (3) if a dominant grass, A. gerardii, utilizes water lifted by other tallgrass prairie species. Broadly, the results shown here suggest that hydraulic lift does not appear to be widespread or common in this system, but isolated instances suggest that this process does occur within tallgrass prairie. The isolated instance of hydraulic lift did not vary by grazing treatment, nor did they result in facilitation for neighboring grasses. We suggest that the topographic complexity of this tallgrass prairie and the high rates of nocturnal transpiration observed in this study likely limit the frequency and occurrence of hydraulic lift. These results suggest that hydraulic lift can be a patchy process, particularly in heterogeneous landscapes.

Savanna Tree Seedlings are Physiologically Tolerant to Nighttime Freeze Events.

Freeze events can be important disturbances in savanna ecosystems, yet the interactive effect of freezing with other environmental drivers on plant functioning is unknown. Here, we investigated physiological responses of South African tree seedlings to interactions of water availability and freezing temperatures. We grew widely distributed South African tree species (Colophospermum mopane, Combretum apiculatum, Acacia nigrescens, and Cassia abbreviata) under well-watered and water-limited conditions and exposed individuals to nighttime freeze events. Of the four species studied here, C. mopane was the most tolerant of lower water availability. However, all species were similarly tolerant to nighttime freezing and recovered within one week following the last freezing event. We also show that water limitation somewhat increased freezing tolerance in one of the species (C. mopane). Therefore, water limitation, but not freezing temperatures, may restrict the distribution of these species, although the interactions of these stressors may have species-specific impacts on plant physiology. Ultimately, we show that unique physiologies can exist among dominant species within communities and that combined stresses may play a currently unidentified role in driving the function of certain species within southern Africa.