Root-To-Shoot Signalling: Assessing The Roles of ‘Up’ In the Up and Down World of Long-Distance Signalling In Planta

Abstract

An important mediator of shoot physiological processes can be the supply of signal molecules (other than water and nutrients) from the root system. Root-to-shoot signalling is often considered to be important in regulating shoot growth and water use when soil conditions change without any demonstrable change in shoot water or nutrient status. Changes in xylem sap composition are often thought to be synonymous with changes in root-to-shoot signalling, even though there is considerable re-cycling of compounds between xylem and phloem. Techniques used to collect xylem sap are reviewed. Elucidating the roles of putative root signal molecules in planta has usually taken priority over identifying the sources of signal molecules in xylem sap. The roles of several signal molecules are considered. This choice is selective, and the failure of known signals to account for observed physiological changes in some systems has lead to the conclusions that other novel signals can be important. The efficacy of a given signal molecule can depend on the shoot water and nutrient status, as demonstrated by variation in stomatal responses to abscisic acid. If such variation is widespread in crop species, this may have implications for the increasing intentional use of root-to-shoot signals to modify crop water use and shoot architecture. Research into root-to-shoot signalling may become increasingly reductionist, in trying to evaluate the contribution of root signals versus local processes to observed physiological changes. However, future challenges are to successfully integrate this basic research into improved crop production systems.