Disentangling the relative impacts of precipitation reduction and vapour pressure deficit (VPD) on plant water dynamics and determining whether acclimation may influence these patterns in the future is an important challenge. Here, we report sap flux density (F ), stomatal conductance (G ), hydraulic conductivity (K ) and xylem anatomy in piñon pine (Pinus edulis) and juniper (Juniperus monosperma) trees subjected to five years of precipitation reduction, atmospheric warming (elevated VPD) and their combined effects. No acclimation occurred under precipitation reduction: lower G and F were found for both species compared to ambient conditions. Warming reduced the sensibility of stomata to VPD for both species but resulted in the maintenance of G and F to ambient levels only for piñon. For juniper, reduced soil moisture under warming negated benefits of stomatal adjustments and resulted in reduced F , G and K . Although reduced stomatal sensitivity to VPD also occurred under combined stresses, reductions in G , F and K took place to similar levels as under single stresses for both species. Our results show that stomatal conductance adjustments to high VPD could minimize but not entirely prevent additive effects of warming and drying on water use and carbon acquisition of trees in semi-arid regions.

We examined the effect of soil microbial communities on plant physiological responses to drought. Bouteloua gracilis seeds were planted in sterilized sand with (inoculated) and without (controls) soil microbial communities. After substantial growth, drought was imposed by completely withholding water. Before soil moisture declined to zero, inoculated plants germinated faster, were significantly taller, and maintained greater soil moisture than controls. The greater soil moisture of the inoculated plants allowed greater photosynthesis but also induced lower tissue drought tolerance (as indicated by turgor loss point) compared to controls. The inoculated plants were more susceptible to severe drought compared to control plants as indicated by significantly lower mean stomatal conductance, as well as marginally significantly greater mean wilting score, for the entire severe drought period after soil moisture declined to zero. Inoculated plants exhibited enhanced growth and photosynthesis and dampened drought stress over short timescales, but also increased susceptibility to drought over long timescales. This work demonstrates (1) an unexpected insight that microbes can have positive initial effects on plant performance, but negative impacts on plant performance during severe drought, and (2) that microbially altered effects on plant function during well-watered and moderate drought conditions can influence plant function under subsequent severe drought.

Climate warming should result in hotter droughts of unprecedented severity in this century. Such droughts have been linked with massive tree mortality, and data suggest that warming interacts with drought to aggravate plant performance. Yet how forests will respond to hotter droughts remains unclear, as does the suite of mechanisms trees use to deal with hot droughts. We used an ecosystem-scale manipulation of precipitation and temperature on piñon pine (Pinus edulis) and juniper (Juniperus monosperma) trees to investigate nitrogen (N) cycling-induced mitigation processes related to hotter droughts. We found that while negative impacts on plant carbon and water balance are manifest after prolonged drought, performance reductions were not amplified by warmer temperatures. Rather, increased temperatures for 5 years stimulated soil N cycling under piñon trees and modified tree N allocation for both species, resulting in mitigation of hotter drought impacts on tree water and carbon functions. These findings suggest that adjustments in N cycling are likely after multi-year warming conditions and that such changes may buffer reductions in tree performance during hotter droughts. The results highlight our incomplete understanding of trees' ability to acclimate to climate change, raising fundamental questions about the resistance potential of forests to long-term, compound climatic stresses.

Plants close their stomata during drought to avoid excessive water loss, but species differ in respect to the drought severity at which stomata close. The stomatal closure point is related to xylem anatomy and vulnerability to embolism, but it also has implications for phloem transport and possibly phloem anatomy to allow sugar transport at low water potentials. Desiccation-tolerant plants that close their stomata at severe drought should have smaller xylem conduits and/or fewer and smaller interconduit pits to reduce vulnerability to embolism but more phloem tissue and larger phloem conduits compared with plants that avoid desiccation. These anatomical differences could be expected to increase in response to long-term reduction in precipitation. To test these hypotheses, we used tridimensional synchroton X-ray microtomograph and light microscope imaging of combined xylem and phloem tissues of 2 coniferous species: one-seed juniper (Juniperus monosperma) and piñon pine (Pinus edulis) subjected to precipitation manipulation treatments. These species show different xylem vulnerability to embolism, contrasting desiccation tolerance, and stomatal closure points. Our results support the hypothesis that desiccation tolerant plants require higher phloem transport capacity than desiccation avoiding plants, but this can be gained through various anatomical adaptations in addition to changing conduit or tissue size.

Understanding how climate alters plant-soil water dynamics, and its impact on physiological functions, is critical to improved predictions of vegetation responses to climate change. Here we analyzed how belowground interactions for water shift under warming and drought, and associated impacts on plant functions. In a semi-arid woodland, adult trees (piñon and juniper) and perennial grasses (blue grama) were exposed to warming and precipitation reduction. After 6 years of continuous treatment exposure, soil and plant water isotopic composition was measured to assess plant water uptake depths and community-level water source partitioning. Warming and drought modified plant water uptake depths. Under warming, contrasting changes in water sources between grasses and trees reduced belowground water source partitioning, resulting in higher interspecific competition for water. Under drought, shifts in trees and grass water sources to deeper soil layers resulted in the maintenance of the naturally occurring water source partitioning among species. Trees showed higher water stress, and reduced water use and photosynthesis in response to warming and drought. This case study demonstrates that neighboring plants shift their competitive interactions for water under prolonged warming and drought, but regardless of whether changes in moisture sources will result in increased competition among species or maintained partitioning of water resources, these competitive adaptations may easily be overridden by climate extremes.