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Plant survival depends on a balance between carbon supply and demand. When carbon supply becomes limited, plants buffer demand by using stored carbohydrates (sugar and starch). During drought, NSCs (non-structural carbohydrates) may accumulate if growth stops before photosynthesis. This expectation is pervasive, yet few studies have combined simultaneous measurements of drought, photosynthesis, growth, and carbon storage to test this. Using a field experiment with mature trees in a semi-arid woodland, we show that growth and photosynthesis slow in parallel as$${\psi }_{{pd}}$$${\psi }_{pd}$declines, preventing carbon storage in two species of conifer (J. monospermaandP. edulis). During experimental drought, growth and photosynthesis were frequently co-limited. Our results point to an alternative perspective on how plants use carbon that views growth and photosynthesis as independent processes both regulated by water availability.

Non‐structural carbohydrates (NSCs), as the labile fraction and dominant carbon currency, are essential mediators of plant adaptation to environments. However, whether and how NSC coordinates with plant economic strategy frameworks, particularly the well‐recognized leaf economics spectrums (LES) and root economics space (RES), remains unclear.

Frequent observations of higher mortality in larger trees than in smaller ones during droughts have sparked an increasing interest in size‐dependent drought‐induced mortality. However, the underlying physiological mechanisms are not well understood, with height‐associated hydraulic constraints often being implied as the potential mechanism driving increased drought vulnerability. We performed a quantitative synthesis on how key traits that drive plant water and carbon economy change with tree height within species and assessed the implications that the different constraints and compensations may have on the interacting mechanisms (hydraulic failure, carbon starvation and/or biotic‐agent attacks) affecting tree vulnerability to drought. While xylem tension increases with tree height, taller trees present a range of structural and functional adjustments, including more efficient water use and transport and greater water uptake and storage capacity, that mitigate the path‐length‐associated drop in water potential. These adaptations allow taller trees to withstand episodic water stress. Conclusive evidence for height‐dependent increased vulnerability to hydraulic failure and carbon starvation, and their coupling to defence mechanisms and pest and pathogen dynamics, is still lacking. Further research is needed, particularly at the intraspecific level, to ascertain the specific conditions and thresholds above which height hinders tree survival under drought.

Within vascular plants, the partitioning of hydraulic resistance along the soil‐to‐leaf continuum affects transpiration and its response to environmental conditions. In trees, the fractional contribution of leaf hydraulic resistance (R_leaf) to total soil‐to‐leaf hydraulic resistance (R_total), or fR_leaf(=R_leaf/R_total), is thought to be large, but this has not been tested comprehensively. We compiled a multibiome data set of fR_leafusing new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fR_leafaveraged 0.51 (95% confidence interval [CI] = 0.46−0.57) and it declined with tree height. We also used the allometric relationship between field‐based measurements of soil‐to‐leaf hydraulic conductance and laboratory‐based measurements of leaf hydraulic conductance to compute the average fR_leaffor 19 tree samples, which was 0.40 (95% CI = 0.29−0.56). The in situ technique produces a more accurate descriptor of fR_leafbecause it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fR_leafmay help stems from loss of hydraulic conductance. Thus, the decline in fR_leafwith tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality.

Heat and drought affect plant chemical defenses and thereby plant susceptibility to pests and pathogens. Monoterpenes are of particular importance for conifers as they play critical roles in defense against bark beetles. To date, work seeking to understand the impacts of heat and drought on monoterpenes has primarily focused on young potted seedlings, leaving it unclear how older age classes that are more vulnerable to bark beetles might respond to stress. Furthermore, we lack a clear picture of what carbon resources might be prioritized to support monoterpene synthesis under drought stress. To address this, we measured needle and woody tissue monoterpene concentrations and physiological variables simultaneously from mature piñon pines (Pinus edulis) from a unique temperature and drought manipulation field experiment. While heat had no effect on total monoterpene concentrations, trees under combined heat and drought stress exhibited ~ 85% and 35% increases in needle and woody tissue, respectively, over multiple years. Plant physiological variables like maximum photosynthesis each explained less than 10% of the variation in total monoterpenes for both tissue types while starch and glucose + fructose measured 1-month prior explained ~ 45% and 60% of the variation in woody tissue total monoterpene concentrations. Although total monoterpenes increased under combined stress, some key monoterpenes with known roles in bark beetle ecology decreased. These shifts may make trees more favorable for bark beetle attack rather than well defended, which one might conclude if only considering total monoterpene concentrations. Our results point to cumulative and synergistic effects of heat and drought that may reprioritize carbon allocation of specific non-structural carbohydrates toward defense.

Shifts in the age or turnover time of non‐structural carbohydrates (NSC) may underlie changes in tree growth under long‐term increases in drought stress associated with climate change. But NSC responses to drought are challenging to quantify, due in part to large NSC stores in trees and subsequently long response times of NSC to climate variation.