Unravelling response-specificity in Ca²⁺ signalling pathways in plant cells

Considerable advances have been made, both in the technologies available to study changes in intracellular cytosolic free Ca²⁺ ([Ca²⁺ ]_i ), and in our understanding of Ca²⁺ signalling cascades in plant cells, but how specificity can be generated from such a ubiquitous component as Ca²⁺ is questionable. Recently the concept of 'Ca²⁺ signatures' has been formulated; tight control of the temporal and spatial characteristics of alterations in [Ca²⁺ ]_i signals is thought to be responsible, at least in part, for the specificity of the response. However, the way in which Ca²⁺ signatures are decoded, which depends on the nature and location of the targets of the Ca²⁺ signals, has received little attention. In a few key systems, progress is being made on how diverse Ca²⁺ signatures might be transduced within cells in response to specific signals. Valuable pieces of the signal-specificity puzzle are being put together and this is illustrated here using some key examples; these emphasize the global importance of Ca²⁺ -mediated signal-transduction cascades in the responses of plants to a wide diversity of extracellular signals. However, the way in which signal specificity is encoded and transduced is still far from clear. Contents Summary 7 I. Introduction: Ca²⁺ as a signal transducer 8 II. Alterations in intracellular [Ca²⁺ ] 8 1. Measuring alterations in [Ca²⁺ ] 8 Imaging [Ca²⁺ ]_i using Ca²⁺ -sensitive dyes 8 Measuring [Ca²⁺ ]_i using aequorin 9 Imaging [Ca²⁺ ]_i using cameleon 10 2. The concept of the 'Ca²⁺ signature 10 3. How might specific Ca²⁺ signatures be generated? 11 Control of intracellular Ca²⁺ release 11 Control of influx of extracellular Ca²⁺ 12 4. Examples of Ca²⁺ signatures and cellular responses to increases in [Ca²⁺ ] 13 Ca²⁺ signatures in stomatal guard cells in response to abscisic acid signals 14 Ca²⁺ signals in response to abiotic stimuli1 8 Ca²⁺ signatures involved in plant-pathogen responses 19 Ca²⁺ signatures in control of plant reproduction 20 Ca²⁺ signatures in root hairs in response to nodulation signals 23 III. Decoding the [Ca²⁺ ]_i signatures 24 1. Coupling Ca²⁺ signals to responses through CaM 26 2. Coupling Ca²⁺ signals to responses through CDPK 27 3. Novel Ca²⁺ binding proteins as primary Ca²⁺ sensors 28 Conclusions and Perspective 28 References 29.