Electrocyte physiology: 50 years later.

Weakly electric gymnotiform and mormyrid fish generate and detect weak electric fields to image their worlds and communicate. These multi-purpose electric signals are generated by electrocytes, the specialized electric organ (EO) cells that produce the electric organ discharge (EOD). Just over 50 years ago the first experimental analyses of electrocyte physiology demonstrated that the EOD is produced and shaped by the timing and waveform of electrocyte action potentials (APs). Electrocytes of some species generate a single AP from a distinct region of excitable membrane, and this AP waveform determines EOD waveform. In other species, electrocytes possess two independent regions of excitable membrane that generate asynchronous APs with different waveforms, thereby increasing EOD complexity. Signal complexity is further enhanced in some gymnotiforms by the spatio-temporal activation of distinct EO regions with different electrocyte properties. For many mormyrids, additional EOD waveform components are produced by APs that propagate along stalks that connect postsynaptic regions to the main body of the electrocyte. I review here the history of research on electrocyte physiology in weakly electric fish, as well as recent discoveries of key phenomena not anticipated during early work in this field. Recent areas of investigation include the regulation of electrocyte activity by steroid and peptide hormones, the molecular evolution of electrocyte ion channels, and the evolutionary selection of ion channels expressed in excitable cells. These emerging research areas have generated renewed interest in electrocyte function and clear future directions for research addressing a broad range of new and important questions.