REVIEW ARTICLE Stay wet or else: three ways in which plants can adjust hydraulically to their environment

The literature on whole-plant acclimation to drought is reviewed and it is proposed that leaf-level homeostasis in water status is attained during ontogeny largely thanks to whole-plant changes in physical resistance to liquid water flow caused by morphological and anatomical adjustments. It is shown that, in response to water deficits, plant resistance changes at different levels (tissue, organ, individual), levels that are corre-lated with the time scale of the response. It was found that such adjustments apparently tend to increase resistance to flow in the short term and to reduce it in the long term. A critical view of those findings is provided based on the principle that drought-induced changes cannot be analysed separately from the allo-metric changes that take place through ontogeny, as for example proposed by the widely cited hydraulic limita-tion hypothesis. A graphic synthetic model is presented according to which developmental responses to water deficits operate largely through reductions in whole-plant water transport capacity, combined with more or less strong reductions in leaf area (different ‘hydraulic allometries’), depending on the intrinsic tolerance of leaf tissues to partial desiccation. The model is used to show that, as the result of such adjustments, the water transport capacity per unit leaf area can decrease, remain constant, or increase, and it is argued that the expected leaf-level response would be different in each case, respectively involving a decreased, constant, or increased potential for transpiration. The article ends with a plea to collect the evidence needed to evaluate the occurrence of these three different response types across taxa and their association with different environ-ments, including the reanalysis of existing data. Key words: Allometry, drought, hydraulic architecture, hydraulic resistance, ontogeny. ‘Maximizing gas exchange while avoiding hydraulic failure means operating on the edge of dysfunction’ John S. Sperry (2004