Long-distance signalling from roots to shoots assessed: the flooding story.

Several kinds of signal may be generated when roots are exposed to an environmental stress. Some, but not all, are conveyed to shoots in the transpiration stream. Principles are summarized that may help establish experimentally the presence or intensity of root signals transported by transpiration. In many dryland species, flooding of the soil induces developmental responses in the shoot such as epinastic leaf curvature, stomatal closure and slowing of leaf expansion. These reactions compensate for diminished input of resources from the roots. They lend themselves to the study of root-to-shoot signalling by commencing after a time lag of only a few hours, by persisting for several days and by being highly reproducible. Evidence implicating chemical and hydraulic signals in promoting stomatal closure and epinastic curvature in flooded plants is reviewed. Further progress will depend upon examining a wider range of putative signals, accounting for any interactions between them and improving methods for the evaluation of signal durability in transit, and effectiveness at target sites.