• Energy Research

AbstractHigher temperatures associated with climate change are anticipated to trigger an earlier start to the growing season, which could increase the terrestrial C sink strength. Greater variability in the amount and timing of precipitation is also expected with higher temperatures, bringing increased drought stress to many ecosystems. We experimentally assessed the effects of higher temperature and drought on the foliar phenology and shoot growth of mature trees of two semiarid conifer species. We exposed field‐grown trees to a ~45% reduction in precipitation with a rain‐out structure (‘drought’), a ~4.8 °C temperature increase with open‐top chambers (‘heat’), and a combination of both simultaneously (‘drought + heat’). Over the 2013 growing season, drought, heat, and drought + heat treatments reduced shoot and needle growth in piñon pine (Pinus edulis) by ≥39%, while juniper (Juniperus monosperma) had low growth and little response to these treatments. Needle emergence on primary axis branches of piñon pine was delayed in heat, drought, and drought + heat treatments by 19–57 days, while secondary axis branches were less likely to produce needles in the heat treatment, and produced no needles at all in the drought + heat treatment. Growth of shoots and needles, and the timing of needle emergence correlated inversely with xylem water tension and positively with nonstructural carbohydrate concentrations. Our findings demonstrate the potential for delayed phenological development and reduced growth with higher temperatures and drought in tree species that are vulnerable to drought and reveal potential mechanistic links to physiological stress responses. Climate change projections of an earlier and longer growing season with higher temperatures, and consequent increases in terrestrial C sink strength, may be incorrect for regions where plants will face increased drought stress with climate change.

Summary The persistence of vegetation under climate change will depend on a plant's capacity to exploit water resources. We analyzed water source dynamics in piñon pine and juniper trees subjected to precipitation reduction, atmospheric warming, and to both simultaneously. Piñon and juniper exhibited different and opposite shifts in water uptake depth in response to experimental stress and background climate over 3 yr. During a dry summer, juniper responded to warming with a shift to shallow water sources, whereas piñon pine responded to precipitation reduction with a shift to deeper sources in autumn. In normal and wet summers, both species responded to precipitation reduction, but juniper increased deep water uptake and piñon increased shallow water uptake. Shifts in the utilization of water sources were associated with reduced stomatal conductance and photosynthesis, suggesting that belowground compensation in response to warming and water reduction did not alleviate stress impacts for gas exchange. We have demonstrated that predicted climate change could modify water sources of trees. Warming impairs juniper uptake of deep sources during extended dry periods. Precipitation reduction alters the uptake of shallow sources following extended droughts for piñon. Shifts in water sources may not compensate for climate change impacts on tree physiology.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

AbstractClimate 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.