Despite decades of research on plant drought tolerance, the physiological mechanisms by which trees succumb to drought are still under debate. We report results from an experiment designed to separate and test the current leading hypotheses of tree mortality. We show that piñon pine (Pinus edulis) trees can die of both hydraulic failure and carbon starvation, and that during drought, the loss of conductivity and carbohydrate reserves can also co‐occur. Hydraulic constraints on plant carbohydrate use determined survival time: turgor loss in the phloem limited access to carbohydrate reserves, but hydraulic control of respiration prolonged survival. Our data also demonstrate that hydraulic failure may be associated with loss of adequate tissue carbohydrate content required for osmoregulation, which then promotes failure to maintain hydraulic integrity.

Drought challenges plant water uptake and the vascular system. In the xylem it causes embolism that impairs water transport from the soil to the leaves and, if uncontrolled, may even lead to plant mortality via hydraulic failure. What happens in the phloem, however, is less clear because measuring phloem transport is still a significant challenge to plant science. In all vascular plants, phloem and xylem tissues are located next to each other, and there is clear evidence that these tissues exchange water. Therefore, drought should also lead to water shortage in the phloem. In this review, theories used in phloem transport models have been applied to drought conditions, with the goal of shedding light on how phloem transport failure might occur. The review revealed that phloem failure could occur either because of viscosity build-up at the source sites or by a failure to maintain phloem water status and cell turgor. Which one of these dominates depends on the hydraulic permeability of phloem conduit walls. Impermeable walls will lead to viscosity build-up affecting flow rates, while permeable walls make the plant more susceptible to phloem turgor failure. Current empirical evidence suggests that phloem failure resulting from phloem turgor collapse is the more likely mechanism at least in relatively isohydric plants.

Bubble formation in the conduits of woody plants sets a challenge for uninterrupted water transportation from the soil up to the canopy. Freezing and thawing of stems has been shown to increase the number of air-filled (embolized) conduits, especially in trees with large conduit diameters. Despite numerous experimental studies, the mechanisms leading to bubble formation during freezing have not been addressed theoretically. We used classical nucleation theory and fluid mechanics to show which mechanisms are most likely to be responsible for bubble formation during freezing and what parameters determine the likelihood of the process. Our results confirm the common assumption that bubble formation during freezing is most likely due to gas segregation by ice. If xylem conduit walls are not permeable to the salts expelled by ice during the freezing process, osmotic pressures high enough for air seeding could be created. The build-up rate of segregated solutes in front of the ice-water interface depends equally on conduit diameter and freezing velocity. Therefore, bubble formation probability depends on these variables. The dependence of bubble formation probability on freezing velocity means that the experimental results obtained for cavitation threshold conduit diameters during freeze/thaw cycles depend on the experimental setup; namely sample size and cooling rate. The velocity dependence also suggests that to avoid bubble formation during freezing trees should have narrow conduits where freezing is likely to be fast (e.g., branches or outermost layer of the xylem). Avoidance of bubble formation during freezing could thus be one piece of the explanation why xylem conduit size of temperate and boreal zone trees varies quite systematically.