Lack of vulnerability segmentation in four angiosperm tree species: evidence from direct X-ray microtomography observation

KEY MESSAGE : Xylem vulnerability to drought-induced embolism did not differ between stems and petioles of four woody species (Betula pendula , Liriodendron tulipifera , Populus tremula and Olea europaea). Our results, together with data compiled from published literature, indicate that hydraulic segmentation during drought stress is not consistently driven by difference in vulnerability to embolism between stem and terminal organs. CONTEXT: Hydraulic failure and disconnection of distal organs during protracted drought stress is thought to protect large branches or trunks by reducing water loss and restricting the spread of embolism. Hydraulic segmentation and preferential sacrifice of distal organs such as leaves can be driven by two mechanisms: more negative water potentials at the terminal section of the hydraulic pathway and/or by higher vulnerability to xylem embolism of distal organs. Although vulnerability segmentation has been reported in the literature, the generality of this phenomenon is unclear, in part due to the methodological limitations related to direct measurement of xylem vulnerability to embolism in intact plants. AIMS: The objective of this study was to evaluate vulnerability segmentation between petioles and stems using non-invasive micro computed tomography (microCT). METHODS: Vulnerability to embolism was measured in leaf petioles and subtending stems of four woody species (Betula pendula R., Liriodendron tulipifera L., Populus tremula L. and Olea europaea L.) with contrasting drought tolerances. In addition, previously published vulnerability data for petioles and stems were compiled from the literature to investigate the commonality of hydraulic segmentation across a wide range of woody species, with the vulnerability curve methodology distinguished. RESULTS: Using non-invasive imaging on intact plants, we found no evidence of hydraulic segmentation between petioles and stems of four angiosperm tree species, regardless of mechanism. Moreover, the literature dataset indicated that little or no difference in vulnerability to embolism is present between petioles and stems when vulnerability curves were constructed using methods specifically measuring the dynamics of xylem tissue during dehydration (e.g. optical visualization, MicroCT). CONCLUSION: Our results suggest that vulnerability segmentation between stems and distal organs (petioles and leaves) is limited when only xylem tissue is considered. Large differences in vulnerability between stems and leaves are likely to be driven by extra-xylary components, rather than xylem embolism.