Mechanical signalling, calcium and plant form

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Abstract
Calcium is a dynamic signalling molecule which acts to transduce numerous signals in plant tissues. The basis of calcium signalling is outlined and the necessity for measuring and imaging of calcium indicated. Using plants genetically transformed with a cDNA for the calcium-sensitive luminescent protein, aequorin, we have shown touch and wind signals to immediately increase cytosol calcium. Touch and wind signal plant cells mechanically, through tension and compression of appropiate cells. Many plant tissues and cells are very sensitive to mechanical stimulation and the obvious examples of climbing plants, insectivorous species as well as other less well-known examples are described. Touch sensing in these plants may be a simple evolutionary modification of sensitive mechanosensing system present in every plant. The possibility that gravitropism may be a specific adaptation of touch sensing is discussed. There is a growing appreciation that plant form may have a mechanical basis. A simple mechanical mechanism specifying spherical, cylindrical and flat-bladed structures is suggested. The limited morphological variety of plant tissues may also reflect mechanical specification. The article concludes with a discussion of the mechanisms of mechanical sensing, identifying integrin-like molecules as one important component, and considers the specific role of calcium.