Voltage-Dependent Events in the Dendritic Tree
So far, we worked under the convenient fiction that active, voltage-dependent membrane conductances are confined to the spike initiation zone at or close to the cell body and that the dendritic tree is essentially passive. Under the influence of one-dimensional passive cable theory, as refined by Rail and his school (Chaps. 2 and 3), the passive model of dendritic integration of synaptic inputs has become dominant and is taught in all the textbooks. Paradoxically, from the earliest days of intracellular recordings from the fat dendrites of spinal cord motoneurons with the aid of glass microelectrodes, active dendritic responses had been witnessed (Brock, Coombs, and Eccles, 1952; Eccles, Libet, and Young, 1958). Today, there exists overwhelming evidence for a host of voltage-dependent sodium and calcium-conductances in the dendritic tree. In the following section we summarize the experimental evidence and discuss current biophysical modeling efforts focusing on the question of the existence and genesis of fast all-or-none electrical events in the dendrites. We then turn toward possible functional roles of active dendritic processing. One word of advice. It has been argued that linear cable theory as applied to dendrites and taught in the first chapters of this book is irrelevant in the face of all this evidence for active processing and can be relegated to the dustbin. However, this would be a mistake. Under many physiological conditions these nonlinearities will not be relevant. Even if they are, the resistive and capacitive cable properties of the dendrites profoundly influence the initiation and propagation of dendritic action potentials and other active phenomena. Thus, for a complete understanding of the events in active dendritic trees we need to be thoroughly versed in cable theory. The issue of dendritic all-or-none electrical events must be seen as separate from the broader question of the existence and nature of active, that is, voltage-dependent, membrane conductances in the dendritic tree.