Shower thoughts: why scientists should spend more time in the rain.

Stormwater is a vital resource and dynamic driver of terrestrial ecosystem processes. However, processes controlling interactions during and shortly after storms are often poorly seen and poorly sensed when direct observations are substituted with technological ones. We discuss how human observations complement technological ones and the benefits of scientists spending more time in the storm. Human observation can reveal ephemeral storm-related phenomena such as biogeochemical hot moments, organismal responses, and sedimentary processes that can then be explored in greater resolution using sensors and virtual experiments. Storm-related phenomena trigger lasting, oversized impacts on hydrologic and biogeochemical processes, organismal traits or functions, and ecosystem services at all scales. We provide examples of phenomena in forests, across disciplines and scales, that have been overlooked in past research to inspire mindful, holistic observation of ecosystems during storms. We conclude that technological observations alone are insufficient to trace the process complexity and unpredictability of fleeting biogeochemical or ecological events without the shower thoughts produced by scientists' human sensory and cognitive systems during storms.

Leaf habit affects the distribution of drought sensitivity but not water transport efficiency in the tropics.

Considering the global intensification of aridity in tropical biomes due to climate change, we need to understand what shapes the distribution of drought sensitivity in tropical plants. We conducted a pantropical data synthesis representing 1117 species to test whether xylem-specific hydraulic conductivity (KS ), water potential at leaf turgor loss (ΨTLP ) and water potential at 50% loss of KS (ΨP50 ) varied along climate gradients. The ΨTLP and ΨP50 increased with climatic moisture only for evergreen species, but KS did not. Species with high ΨTLP and ΨP50 values were associated with both dry and wet environments. However, drought-deciduous species showed high ΨTLP and ΨP50 values regardless of water availability, whereas evergreen species only in wet environments. All three traits showed a weak phylogenetic signal and a short half-life. These results suggest strong environmental controls on trait variance, which in turn is modulated by leaf habit along climatic moisture gradients in the tropics.

Quantifying and manipulating the angles of light in experimental measurements of plant gas exchange.

Diffuse light has been shown to alter plant leaf photosynthesis, transpiration and water-use efficiency. Despite this, the angular distribution of light for the artificial light sources used with common gas exchange systems is unknown. Here, we quantify the angular distribution of light from common gas exchange systems and demonstrate the use of an integrating sphere for manipulating those light distributions. Among three different systems, light from a 90° angle perpendicular to the leaf surface (±5.75°) was <25% of the total light reaching the leaf surface. The integrating sphere resulted in a greater range of possible distributions from predominantly direct light (i.e., >40% of light from a 90 ± 5.75° angle perpendicular to the leaf surface) to almost entirely diffuse (i.e., light from an even distribution drawn from a nearly 0° horizontal angle to a perpendicular 90° angle). The integrating sphere can thus create light environments that more closely mimic the variation in sunlight under both clear and cloudy conditions. In turn, different proportions of diffuse light increased, decreased or did not change photosynthetic rates depending on the plant species observed. This new tool should allow the scientific community to explore new and creative questions about plant function within the context of global climate change.

Diffuse light and wetting differentially affect tropical tree leaf photosynthesis.

Most ecosystems experience frequent cloud cover resulting in light that is predominantly diffuse rather than direct. Moreover, these cloudy conditions are often accompanied by rain that results in wet leaf surfaces. Despite this, our understanding of photosynthesis is built upon measurements made on dry leaves experiencing direct light. Using a modified gas exchange setup, we measured the effects of diffuse light and leaf wetting on photosynthesis in canopy species from a tropical montane cloud forest. We demonstrate significant variation in species-level response to light quality independent of light intensity. Some species demonstrated 100% higher rates of photosynthesis in diffuse light, and others had 15% greater photosynthesis in direct light. Even at lower light intensities, diffuse light photosynthesis was equal to that under direct light conditions. Leaf wetting generally led to decreased photosynthesis, particularly when the leaf surface with stomata became wet; however, there was significant variation across species. Ultimately, we demonstrate that ecosystem photosynthesis is significantly altered in response to environmental conditions that are ubiquitous. Our results help to explain the observation that net ecosystem exchange can increase in cloudy conditions and can improve the representation of these processes in Earth systems models under projected scenarios of global climate change.

Spatiotemporal Coupling of Vessel Cavitation and Discharge of Stored Xylem Water in a Tree Sapling.

Water discharge from stem internal storage compartments is thought to minimize the risk of vessel cavitation. Based on this concept, one would expect that water storage compartments involved in the buffering of xylem tensions empty before the onset of vessel cavitation under drought stress, and potentially refill after soil saturation. However, scant in vivo data exist that elucidate this localized spatiotemporal coupling. In this study on intact saplings of American chestnut (Castanea dentata), x-ray computed microtomography (microCT) showed that the xylem matrix surrounding vessels releases stored water and becomes air-filled either concurrent to or after vessel cavitation under progressive drought stress. Among annual growth rings, the xylem matrix of the current year remained largely water-filled even under severe drought stress. In comparison, microtomography images collected on excised stems showed that applied pressures of much greater than 0 MPa were required to induce water release from the xylem matrix. Viability staining highlighted that water release from the xylem matrix was associated primarily with emptying of dead fibers. Refilling of the xylem matrix and vessels was detected in intact saplings when the canopy was bagged and stem water potential was close to 0 MPa, and in leafless saplings over the winter period. In conclusion, this study indicates that the bulk of water stored in the xylem matrix is released after the onset of vessel cavitation, and suggests that capillary water contributes to overall stem water storage under drought but is not used primarily for the prevention of drought-induced vessel cavitation in this species.

Foliar water uptake: Processes, pathways, and integration into plant water budgets.

Nearly all plant families, represented across most major biomes, absorb water directly through their leaves. This phenomenon is commonly referred to as foliar water uptake. Recent studies have suggested that foliar water uptake provides a significant water subsidy that can influence both plant water and carbon balance across multiple spatial and temporal scales. Despite this, our mechanistic understanding of when, where, how, and to what end water is absorbed through leaf surfaces remains limited. We first review the evidence for the biophysical conditions necessary for foliar water uptake to occur, focusing on the plant and atmospheric water potentials necessary to create a gradient for water flow. We then consider the different pathways for uptake, as well as the potential fates of the water once inside the leaf. Given that one fate of water from foliar uptake is to increase leaf water potentials and contribute to the demands of transpiration, we also provide a quantitative synthesis of observed rates of change in leaf water potential and total fluxes of water into the leaf. Finally, we identify critical research themes that should be addressed to effectively incorporate foliar water uptake into traditional frameworks of plant water movement.

Linking coordinated hydraulic traits to drought and recovery responses in a tropical montane cloud forest.

PREMISE: Understanding plant hydraulic functioning and water balance during drought has become key in predicting species survival and recovery. However, there are few insightful studies that couple physiological and morphological attributes for many ecosystems, such as the vulnerable Tropical Montane Cloud Forests (TMCF). In this study, we evaluated drought resistance and recovery for saplings for five tree species spanning deciduous to evergreen habits from a Mexican TMCF. METHODS: In drought simulations, water was withheld until plants reached species-specific P50 or P88 values (pressures required to induce a 50 or 88% loss in hydraulic conductivity), then they were rewatered. Drought resistance was considered within the isohydric-anisohydric framework and compared to leaf gas exchange, water status, pressure-volume curves, specific leaf area, and stomatal density. RESULTS: The TMCF species closed stomata well before significant losses in hydraulic conductivity (isohydric). Yet, despite the coordination of these traits, the traits were not useful for predicting the time needed for the species to reach critical hydraulic thresholds. Instead, maximum photosynthetic rates explained these times, reinforcing the linkage between hydraulic and carbon dynamics. Despite their varying hydraulic conductivities, stomatal responses, and times to hydraulic thresholds, 58 of the 60 study plants recovered after the rewatering. The recovery of photosynthesis and stomatal conductance can be explained by the P50 values and isohydry. CONCLUSIONS: This study raises new questions surrounding drought management strategies, recovery processes, and how lethal thresholds are defined. Further studies need to consider the role of water and carbon balance in allowing for both survival and recovery from drought.

Cloud immersion: an important water source for spruce and fir saplings in the southern Appalachian Mountains.

Cloud immersion can provide a potentially important moisture subsidy to plants in areas of frequent fog including the threatened spruce-fir communities of the southern Appalachian Mountains (USA). These mountaintop communities grow only above ~1,500 m elevation, harbor the endemic Abies fraseri, and have been proposed to exist because of frequent cloud immersion. While several studies have demonstrated the importance of cloud immersion to plant water balance, no study has evaluated the proportion of plant water derived from cloud moisture in this ecosystem. Using the isotopic mixing model, IsoSource, we analyzed the isotopic composition of hydrogen and oxygen for water extracted from ground water, deep soil, shallow soil, fog, and plant xylem at the upper and lower elevational limits both in May (beginning of the growing season) and October (end of the growing season). Cloud-immersion water contributed up to 31% of plant water at the upper elevation sites in May. High-elevation plants of both species also experienced greater cloud immersion and had greater cloud water absorption (14-31%) compared to low-elevation plants (4-17%). Greater cloud water uptake occurred in May compared to October, despite similar rainfall and cloud-immersion frequencies. These results demonstrate the important water subsidy that cloud-immersion water can provide. With a warming climate leading potentially to increases in the ceiling of the cloud base and, thus, less frequent cloud immersion, persistence of these relic mountaintop forests may depend on the magnitude of these changes and the compensating capabilities of other water sources.

Ecophysiological importance of cloud immersion in a relic spruce-fir forest at elevational limits, southern Appalachian Mountains, USA.

Climate warming predicts changes to the frequency and height of cloud-immersion events in mountain communities. Threatened southern Appalachian spruce-fir forests have been suggested to persist because of frequent periods of cloud immersion. These relic forests exist on only seven mountaintop areas, grow only above ca. 1,500 m elevation (maximum 2,037 m), and harbor the endemic Abies fraseri. To predict future distribution, we examined the ecophysiological effects of cloud immersion on saplings of A. fraseri and Picea rubens at their upper and lower elevational limits. Leaf photosynthesis, conductance, transpiration, xylem water potentials, and general abiotic variables were measured simultaneously on individuals at the top (1,960 m) and bottom (1,510 m) of their elevation limits on numerous clear and cloud-immersed days throughout the growing season. The high elevation sites had 1.5 as many cloud-immersed days (75 % of days) as the low elevation sites (56 % of days). Cloud immersion resulted in higher photosynthesis, leaf conductance, and xylem water potentials, particularly during afternoon measurements. Leaf conductance remained higher throughout the day with corresponding increases in photosynthesis and transpiration, despite low photon flux density levels, leading to an increase in water potentials from morning to afternoon. The endemic A. fraseri had a greater response in carbon gain and water balance in response to cloud immersion. Climate models predict warmer temperatures with a decrease in the frequency of cloud immersion for this region, leading to an environment on these peaks similar to elevations where spruce-fir communities currently do not exist. Because spruce-fir communities may rely on cloud immersion for improved carbon gain and water conservation, an upslope shift is likely if cloud ceilings rise. Their ultimate survival will likely depend on the magnitude of changes in cloud regimes.

Tracing plant-environment interactions from organismal to planetary scales using stable isotopes: a mini review.

Natural isotope variation forms a mosaic of isotopically distinct pools across the biosphere and flows between pools integrate plant ecology with global biogeochemical cycling. Carbon, nitrogen, and water isotopic ratios (among others) can be measured in plant tissues, at root and foliar interfaces, and in adjacent atmospheric, water, and soil environments. Natural abundance isotopes provide ecological insight to complement and enhance biogeochemical research, such as understanding the physiological conditions during photosynthetic assimilation (e.g. water stress) or the contribution of unusual plant water or nutrient sources (e.g. fog, foliar deposition). While foundational concepts and methods have endured through four decades of research, technological improvements that enable measurement at fine spatiotemporal scales, of multiple isotopes, and of isotopomers, are advancing the field of stable isotope ecology. For example, isotope studies now benefit from the maturation of field-portable infrared spectroscopy, which allows the exploration of plant-environment sensitivity at physiological timescales. Isotope ecology is also benefiting from, and contributing to, new understanding of the plant-soil-atmosphere system, such as improving the representation of soil carbon pools and turnover in land surface models. At larger Earth-system scales, a maturing global coverage of isotope data and new data from site networks offer exciting synthesis opportunities to merge the insights of single-or multi-isotope analysis with ecosystem and remote sensing data in a data-driven modeling framework, to create geospatial isotope products essential for studies of global environmental change.

Sensitivity and threshold dynamics of Pinus strobus and Quercus spp. in response to experimental and naturally occurring severe droughts.

Increased drought frequency and severity are a pervasive global threat, yet the capacity of mesic temperate forests to maintain resilience in response to drought remains poorly understood. We deployed a throughfall removal experiment to simulate a once in a century drought in New Hampshire, USA, which coupled with the region-wide 2016 drought, intensified moisture stress beyond that experienced in the lifetimes of our study trees. To assess the sensitivity and threshold dynamics of two dominant northeastern tree genera (Quercus and Pinus), we monitored sap flux density (Js), leaf water potential and gas exchange, growth and intrinsic water-use efficiency (iWUE) for one pretreatment year (2015) and two treatment years (2016-17). Results showed that Js in pine (Pinus strobus L.) declined abruptly at a soil moisture threshold of 0.15 m3 m-3, whereas oak's (Quercus rubra L. and Quercus velutina Lam.) threshold was 0.11 m3 m-3-a finding consistent with pine's more isohydric strategy. Nevertheless, once oaks' moisture threshold was surpassed, Js declined abruptly, suggesting that while oaks are well adapted to moderate drought, they are highly susceptible to extreme drought. The radial growth reduction in response to the 2016 drought was more than twice as great for pine as for oaks (50 vs 18%, respectively). Despite relatively high precipitation in 2017, the oaks' growth continued to decline (low recovery), whereas pine showed neutral (treatment) or improved (control) growth. The iWUE increased in 2016 for both treatment and control pines, but only in treatment oaks. Notably, pines exhibited a significant linear relationship between iWUE and precipitation across years, whereas the oaks only showed a response during the driest conditions, further underscoring the different sensitivity thresholds for these species. Our results provide new insights into how interactions between temperate forest tree species' contrasting physiologies and soil moisture thresholds influence their responses and resilience to extreme drought.

Why size matters: the interactive influences of tree diameter distribution and sap flow parameters on upscaled transpiration.

In stands with a broad range of diameters, a small number of very large trees can disproportionately influence stand basal area and transpiration (Et). Sap flow-based Et estimates may be particularly sensitive to large trees due to nonlinear relationships between tree-level water use (Q) and tree diameter at breast height (DBH). Because Q is typically predicted on the basis of DBH and sap flow rates measured in a subset of trees and then summed to obtain Et, we assessed the relative importance of DBH and sap flow variables (sap velocity, Vs, and sapwood depth, Rs) in determining the magnitude of Et and its dependence on large trees in a tropical montane forest ecosystem. Specifically, we developed a data-driven simulation framework to vary the relationship between DBH and Vs and stand DBH distribution and then calculate Q, Et and the proportion of Et contributed by the largest tree in each stand. Our results demonstrate that variation in how Rs is determined in the largest trees can alter estimates up to 26% of Et while variation in how Vs is determined can vary results by up to 132%. Taken together, these results highlight a great need to expand our understanding of water transport in large trees as this hinders our ability to predict water fluxes accurately from stand to catchment scales.

Vegetation-zonation patterns across a temperate mountain cloud forest ecotone are not explained by variation in hydraulic functioning or water relations.

Many studies have demonstrated linkages between the occurrence of fog and ecophysiological functioning in cloud forests, but few have investigated hydraulic functioning as a determining factor that explains sharp changes in vegetation. The objective of this study was to compare the plant water status during cloud-immersed and non-immersed conditions and hydraulic vulnerability in branches and roots of species across a temperate, mountain fog ecotone. Because cloud forests are often dark, cool and very moist, we expected cloud forest species to have less drought-tolerant characteristics (i.e., lower Pe and P50-the pressures required to induce a 12 and 50% loss in hydraulic conductivity, respectively) relative to non-cloud forest species in adjacent (lower elevation) forests. Additionally, due to the ability of cloud forest species to absorb cloud-fog water, we predicted greater improvements in hydraulic functioning during fog in cloud forest species relative to non-cloud forest species. Across the cloud forest ecotone, most species measured were very resistant to losses in conductivity with branch P50 values from -4.5 to -6.0 MPa, hydraulic safety margins (Ψmin - P50) >1.5 MPa and low calculated hydraulic conductivity losses. Roots had greater vulnerabilities, with P50 values ranging from -1.4 to -2.5 MPa, leading to greater predicted losses in conductivity (∼20%). Calculated values suggested strong losses of midday leaf hydraulic conductance in three of the four species, supporting the hydraulic segmentation hypothesis. In both cloud forest and hardwood species, Ψs were greater on foggy days than sunny days, demonstrating the importance of fog periods to plant water balance across fog regimes. Thus, frequent fog did not result in systemic changes in hydraulic functioning or vulnerability to embolism across our temperate cloud forest ecotone. Finally, roots functioned with lower hydraulic conductivity than branches, suggesting that they may serve as more sensitive indicators of hydraulic functioning in these mesic, foggy ecosystems.

Foliar uptake, carbon fluxes and water status are affected by the timing of daily fog in saplings from a threatened cloud forest.

In cloud forests, foliar uptake (FU) of water has been reported for numerous species, possibly acting to relieve daily water and carbon stress. While the prevalence of FU seems common, how daily variation in fog timing may affect this process has not been studied. We examined the quantity of FU, water potentials, gas exchange and abiotic variation at the beginning and end of a 9-day exposure to fog in a glasshouse setting. Saplings of Abies fraseri (Pursh) Poir. and Picea rubens Sarg. were exposed to morning (MF), afternoon (AF) or evening fog (EF) regimes to assess the ability to utilize fog water at different times of day and after sustained exposure to simulated fog. The greatest amount of FU occurred during MF (up to 50%), followed by AF (up to 23%) and then EF, which surprisingly had no FU. There was also a positive relationship between leaf conductance and FU, suggesting a role of stomata in FU. Moreover, MF and AF lead to the greatest improvements in daily water balance and carbon gain, respectively. Foliar uptake was important for improving plant ecophysiology but was influenced by diurnal variation in fog. With climate change scenarios predicting changes to cloud patterns and frequency that will likely alter diurnal patterns, cloud forests that rely on this water subsidy could be affected.