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Xylem resistance to cavitation is an important trait that is related to the ecology and survival of plant species. Vessel network characteristics, such as vessel length and connectivity, could affect the spread of emboli from gas-filled vessels to functional ones, triggering their cavitation. We hypothesized that the cavitation resistance of xylem vessels is randomly distributed throughout the vessel network. We predicted that single vessel air injection (SVAI) vulnerability curves (VCs) would thus be affected by sample length. Longer stem samples were predicted to appear more resistant than shorter samples due to the sampled path including greater numbers of vessels. We evaluated the vessel network characteristics of grapevine (Vitis vinifera L.), English oak (Quercus robur L.) and black cottonwood (Populus trichocarpa Torr. & A. Gray), and constructed SVAI VCs for 5- and 20-cm-long segments. We also constructed VCs with a standard centrifuge method and used computer modelling to estimate the curve shift expected for pathways composed of different numbers of vessels. For all three species, the SVAI VCs for 5 cm segments rose exponentially and were more vulnerable than the 20 cm segments. The 5 cm curve shapes were exponential and were consistent with centrifuge VCs. Modelling data supported the observed SVAI VC shifts, which were related to path length and vessel network characteristics. These results suggest that exponential VCs represent the most realistic curve shape for individual vessel resistance distributions for these species. At the network level, the presence of some vessels with a higher resistance to cavitation may help avoid emboli spread during tissue dehydration.

Many plant lineages, including oaks (Quercus spp.), have both vessels and tracheids as hydraulically conductive cells within their xylem. The structure of these co-occurring conduit types and their contribution to plant hydraulic function have been relatively little studied. We hypothesized that vasicentric tracheids contribute to hydraulic function under conditions of low water availability. We predicted that within a species, oaks growing at drier and warmer low elevation sites would have more tracheids and be more embolism resistant compared to those growing at moister and colder higher elevation sites. We also predicted that across species, lower elevation oaks would have increased tracheid abundance within their xylem. Five oak species differed in many xylem traits, including vessel diameter and length, tracheid size and abundance, embolism resistance, and hydraulic conductivity. Tracheids were most abundant in the xylem of the highest elevation species at sites that receive winter snow and freezing temperatures. Vessels were relatively vulnerable to embolism as confirmed with multiple methods, including centrifuge vulnerability curves, micro-CT scans of native stem samples, and single vessel air injection. Theoretical conductivity calculations indicated that tracheids account for 5.7–15.5% of conductivity in hydrated stems, with tracheids likely increasing in importance as large diameter vulnerable vessels embolize. The occurrence of both vessels and vasicentric tracheids in the xylem of oaks may enable them to function within highly seasonal climates. Tracheids, though often overlooked, may be particularly important in maintaining conductivity throughout much of the year when water potentials decline from seasonal maximums and following freeze-thaw events.

Societal Impact Statement Drought plays a conspicuous role in forest mortality, and is expected to become more severe in future climate scenarios. Recent surges in drought‐associated forest tree mortality have been documented worldwide. For example, recent droughts in California and Texas killed approximately 129 million and 300 million trees, respectively. Drought has also induced acute pine tree mortality across east‐central China, and across extensive areas in southwest China. Understanding the biological processes that enable trees to modify wood development to mitigate the adverse effects of drought will be crucial for the development of successful strategies for future forest management and conservation. Summary Drought is a recurrent stress to forests, causing periodic forest mortality with enormous economic and environmental costs. Wood is the water‐conducting tissue of tree stems, and trees modify wood development to create anatomical features and hydraulic properties that can mitigate drought stress. This modification of wood development can be seen in tree rings where not only the amount of wood but also the morphology of the water‐conducting cells are modified in response to environmental conditions. In this review, we provide an overview of how trees conduct water, and how trees modify wood development to affect water conduction properties in response to drought. We discuss key needs for new research, and how new knowledge of wood formation can play a role in the conservation of forests under threat by climate change.

Xylem vessel structure changes as trees grow and mature. Age‐ and development‐related changes in xylem structure are likely related to changes in hydraulic function. We examined whether hydraulic function, including hydraulic conductivity and vulnerability to water‐stress‐induced xylem embolism, changed over the course of cambial development in the stems of 17 tree species. We compared current‐year growth of young (1–4 years), intermediate (2–7 years), and older (3–10 years) stems occurring in series along branches. Diffuse and ring porous species were examined, but nearly all species produced only diffuse porous xylem in the distal branches that were examined irrespective of their mature xylem porosity type. Vessel diameter and length increased with cambial age. Xylem became both more conductive and more cavitation resistant with cambial age. Ring porous species had longer and wider vessels and xylem that had higher conductivity and was more vulnerable to cavitation; however, these differences between porosity types were not present in young stem samples. Understanding plant hydraulic function and architecture requires the sampling of multiple‐aged tissues because plants may vary considerably in their xylem structural and functional traits throughout the plant body, even over relatively short distances and closely aged tissues.

During secondary growth, forest trees can modify the anatomy of the wood produced by the vascular cambium in response to environmental conditions. Notably, the trees of the model angiosperm genus, Populus, reduce the risk of cavitation and hydraulic failure under water stress by producing water-conducting vessel elements with narrow lumens, which are more numerous and more interconnected with each other. Here, we determined the genetic architecture of vessel traits affecting hydraulic physiology and resilience to water stress. Vessel traits were measured for clonally replicated genotypes of a unique Populus deltoides x nigra population carrying genomically defined insertions and deletions that create gene dosage variation. We found significant phenotypic variation for all traits measured (mean vessel diameter, height-corrected mean vessel diameter, vessel frequency, height-corrected vessel frequency, vessel grouping index, and mean vessel circularity), and that all traits were under genetic control and showed moderate heritability values, ranging from 0.32 to 0.53. Whole-genome scans of correlations between gene dosage and phenotypic traits identified quantitative trait loci for tree height, mean vessel diameter, height-corrected mean vessel diameter, height-corrected vessel frequency, and vessel grouping index. Our results demonstrate that vessel traits affecting hydraulic physiology are under genetic control, and both pleiotropic and trait-specific quantitative trait loci are found for these traits.