Evolution of excitability in lower metazoans

All forms of life exhibit excitability; it is one of the characteristics by which living creatures can be recognized. In this chapter, we examine the different manifestations of excitability exhibited by the Metazoa and show how elements present in the Bacteria come together in the Protozoa, Porifera, Cnidaria, and Ctenophora (see Fig. 1) to form the patterns of excitability known as “behavior.” We consider the role of excitation in fertilized eggs and conducting epithelia, as well as the origins of signaling in nerves and muscles. We describe different forms of all-or-nothing signaling, as well as ways of generating graded responsiveness. This study attempts to provide a practical approach to understanding the limitations of excitable systems. We believe that defining these limits is more useful than glorifying their seemingly endless sophistication. THE NATURE OF EXCITABILITY Excitability Defined Excitability is easy to recognize; less easy to define. We recognize excitability when we see it, by the way an organism responds to an external stimulus. For there to be a response, stimulus and organism must interact and the organism must “receive” the stimulus. Inevitably, the stimulus site, or receptor, and the response site, or effector, will be at different locations even in single cells. Thus, excitability depends on the transmission of signals from receptor to effector. The signals may be chemical and spread by passive diffusion, electrical and spread by the transfer of ionic charge, or mechanical and spread by a physical disturbance. We focus in this section on the links between chemical...