Bursting Cells

Some neurons throughout the animal kingdom respond to an intracellular current injection or to an appropriate sensory stimulus with a stereotypical sequence of two to five fast spikes riding upon a slow depolarizing envelope. The entire event, termed a burst, is over within 10-40 msec and is usually terminated by a profound afterhyperpolarization (ΑΗΡ). Such bursting cells are not a random feature of a certain fraction of all cells but can be identified with specific neuronal subpopulations. What are the mechanisms generating this intrinsic firing pattern and what is its meaning? Bursting cells can easily be distinguished from a cell firing at a high maintained frequency by the fact that bursts will persist even at a low firing frequency. As illustrated by the thalamic relay cell of Fig. 9.4, some cells can switch between a mode in which they predominantly respond to stimuli via single, isolated spikes and one in which bursts are common. Because we believe that bursting constitutes a special manner of signaling important information, we devote a single, albeit small chapter to this topic. In the following, we describe a unique class of cells that frequently signal with bursts, and we touch upon the possible biophysical mechanisms that give rise to bursting. We finish this excursion by focussing on a functional study of bursting cells in the electric fish and speculate about the functional relevance of burst firing. Neocortical cells are frequently classified according to their response to sustained current injections. While these distinctions are not all or none, there is broad agreement for three classes: regular spiking, fast spiking, and intrinsically bursting neurons (Connors, Gutnick, and Prince, 1982; McCormick et al., 1985; Connors and Gutnick, 1990; Agmon and Connors, 1992; Baranyi, Szente, and Woody, 1993; Nuńez, Amzica, and Steriade, 1993; Gutnick and Crill, 1995; Gray and McCormick, 1996). Additional cell classes have been identified (e.g., the chattering cells that fire bursts of spikes with interburst intervals ranging from 15 to 50 msec; Gray and McCormick, 1996), but whether or not they occur widely has not yet been settled. The cells of interest to us are the intrinsically bursting cells.