Input-output functions of mammalian motoneurons.

Our intent in this review was to consider the relationship between the biophysical properties of motoneurons and the mechanisms by which they transduce the synaptic inputs they receive into changes in their firing rates. Our emphasis has been on experimental results obtained over the past twenty years, which have shown that motoneurons are just as complex and interesting as other central neurons. This work has shown that motoneurons are endowed with a rich complement of active dendritic conductances, and flexible control of both somatic and dendritic channels by endogenous neuromodulators. Although this new information requires some revision of the simple view of motoneuron input-output properties that was prevalent in the early 1980's (see sections 2.3 and 2.10), the basic aspects of synaptic transduction by motoneurons can still be captured by a relatively simple input-output model (see section 2.3, equations 1-3). It remains valid to describe motoneuron recruitment as a product of the total synaptic current delivered to the soma, the effective input resistance of the motoneuron and the somatic voltage threshold for spike initiation (equations 1 and 2). However, because of the presence of active channels activated in the subthreshold range, both the delivery of synaptic current and the effective input resistance depend upon membrane potential. In addition, activation of metabotropic receptors by achetylcholine, glutamate, noradrenaline, serotonin, substance P and thyrotropin releasing factor (TRH) can alter the properties of various voltage- and calcium-sensitive channels and thereby affect synaptic current delivery and input resistance. Once motoneurons are activated, their steady-state rate of repetitive discharge is linearly related to the amount of injected or synaptic current reaching the soma (equation 3). However, the slope of this relation, the minimum discharge rate and the threshold current for repetitive discharge are all subject to neuromodulatory control. There are still a number of unresolved issues concerning the control of motoneuron discharge by synaptic inputs. Under dynamic conditions, when synaptic input is rapidly changing, time- and activity-dependent changes in the state of ionic channels will alter both synaptic current delivery to the spike-generating conductances and the relation between synaptic current and discharge rate. There is at present no general quantitative expression for motoneuron input-output properties under dynamic conditions. Even under steady-state conditions, the biophysical mechanisms underlying the transfer of synaptic current from the dendrites to the soma are not well understood, due to the paucity of direct recordings from motoneuron dendrites. It seems likely that resolving these important issues will keep motoneuron afficiandoes well occupied during the next twenty years.