The vulnerability to drought-induced embolism-conduit diameter link: breaching the anatomy-physiology divide

The best explanations of the relationship between organismal form and function-those regarded by scientists as the most solid — always account for both comparative, across-species, patterns, as well as experimental results. This is true in all of biology, as it is for the study of xylem structure-function relations, where there is still a need for xylem physiology and functional comparative wood anatomy to mutually complement each other. To illustrate the magnitude and urgency of this need, we discuss Sherwin Carlquist’s postulate of a link between vulnerability to drought-induced embolism and conduit diameter, summarizing some of the major global patterns of xylem trait variation that are currently explained by postulating a vulnerability–diameter link. These include wider community mean and maximum vessel diameters in wetter vs drier vegetation types; that vessels can be >700 μm in diameter but plants virtually always produce much narrower ones; that dryland plants with wider vessels drop their leaves earlier; wide-to-narrow vessels across growth rings; and the wide vessels of lianas surrounded by narrow vessels. These patterns are global, and we are aware of no anatomical evidence contradicting a vulnerability–diameter link. Despite the pervasiveness of these patterns, many xylem biologists do not regard the patterns as providing guidance for research in functional xylem biology. Instead, proposing and testing hypotheses to account for all of the data — xylem physiology experiments and comparative anatomical patterns in all their complexity and with all of their contradictions — provides the best way forward for the field. This effort requires proposing and testing hypotheses that are consistent with both experimental as well as comparative data. Crucially, it also requires not rejecting the vulnerability–diameter link without providing an alternative explanation that better explains the patterns currently explained by appeal to the link.