Systemic signaling in response to wounding and pathogens.

Plants possess systemic signaling networks that allow the perception of local stresses to be translated into plant-wide responses. Although information can be propagated via a variety of molecules such as hormones and RNAs moving within the bulk flow of the phloem or in the transpiration stream, the vasculature also appears to be a major pathway whereby extremely rapid signals move bi-directionally throughout the plant. In these cases, the movement mechanisms are not dependent on redistribution through bulk flow. For example, self-reinforcing systems based around changes in Ca2+ and reactive oxygen species, coupled to parallel electrical signaling events appear able to generate waves of information that can propagate at hundreds of μm/s. These signals then elicit distant responses that prime the plant for a more effective defense or stress response in unchallenged tissues. Although ion channels, Ca2+ , reactive oxygen species and associated molecular machineries, such as the NADPH oxidases, have been identified as likely important players in this propagation system, the precise nature of these signaling networks remains to be defined. Critically, whether different stimuli are using the same rapid, systemic signaling network, or whether multiple, parallel pathways for signal propagation are operating to trigger specific systemic outputs remains a key open question.