From local to global: CDPKs in systemic defense signaling upon microbial and herbivore attack.

Calcium-dependent protein kinases (CDPKs) are multifunctional proteins in which a calmodulin-like calcium-sensor and a protein kinase effector domain are combined in one molecule. Not surprisingly, CDPKs were primarily recognized as signaling mediators, which perceive rapid intracellular changes of Ca(2+) ion concentration, for example triggered by environmental stress cues, and relay them into specific phosphorylation events to induce further downstream stress responses. In the context of both, plant exposure to biotrophic pathogens-derived signals as well as plant attack by herbivores and wounding, CDPKs were shown to undergo rapid biochemical activation within seconds to minutes after stimulation and to induce local defence-responses including respective changes in gene expression patterns. In addition, CDPK function was correlated with the control of either salicylic acid-mediated or jasmonic acid-mediated phytohormone signaling pathways, mediating long term resistance to either biotrophic bacterial pathogens or herbivores also in distal parts of a plant. It has long been unclear how an individual enzyme can affect both rapid local as well as long-term distal immune responses. Here, we discuss recently raised topics from the field of CDPK research, in particular with a view on the identification of in vivo phosphorylation targets, which provide first mechanistic insights into the dual role of these enzymes: On the one hand as component of a self-activating circuit responsible for rapid plasma-membrane anchored cell-to-cell signal propagation from local to distal plant sites. On the other hand as nuclear-located regulators of transcription factor activity. Finally, we will highlight the dual function of calcium sensors in plasma-membrane/calcium-mediated signal propagation and in phytohormone signaling-dependent systemic resistance in immune responses to both, bacterial pathogens and herbivores.