Dendritic location of synapses and possible mechanisms for the monosynaptic EPSP in motoneurons.

THE PURPOSE OF THIS PAPER is to bring together recent experimental and theoretical results which, when considered together, add to our understanding of the monosynaptic excitatory postsynaptic potential (EPSP) in moto-neurons. A general mathematical neuron model has been proposed which provides a means of computing EPSP properties that result when synapses are distributed over the soma and dendrites of a nerve cell (29). This general model contains a set of theoretical parameters corresponding to such neu-ronal properties as the membrane time constan .t, the dendritic electro tonic length, the surface area ratio and the condu .ctance ra tio of dendrites to soma, the spatial distribution of synaptic input over the soma .-dendritic surface, and the time course of synaptic current or synaptic conductance. Every specific application of this general model depends upon selecting a set of values for these parameters. We wish to explore the application of this general model to the particular case of the cat spinal motoneuron. Intracellular recordings from motoneurons have provided a description of many of the properties of the monosynaptic EPSP's of these cells (4, 7, 8, 24, 32). In particular, the shapes of the unitary, miniature EPSP's resulting from activation of single Ia afferent fibers have been described. The relationship of these unitary EPSP's to the composite EPSP's elicited by electrical stimulation of many Ia fibers has been discussed (4). We wish to determine whether the experimental data will allow us to put some restrictions on the model parameters which determine EPSP shapes and to provide a more sharply defined picture of the organizati on of synapses on the motoneuronal soma and dendritic tree. In PART I of this paper we will show that a number of restrictions do emerge from\ the analysis. For example, we have found that simple variations in the time course of synaptic currents cannot reproduce the observed EPSP shapes if the synaptic input is distributed uniformly over the motoneuron surface. The most restrictive model which successfully simulates the observed data, and the model which we will discuss in some detail, involves a single time course for the synaptic current, but with the synaptic input occurring in different combinations of somatic and dendritic locations. Thus, a

[1]  P. G. Nelson,et al.  Synaptic Activity in Motoneurons during Natural Stimulation of Muscle Spindles , 1966, Science.

[2]  J. Schadé ON THE VOLUME AND SURFACE AREA OF SPINAL NEURONS. , 1964, Progress in Brain Research.

[3]  Kisou Kubota,et al.  Studies of the Integrative Function of the Motor Neurone , 1963 .

[4]  M. V. Bennett,et al.  Electrotonic Junctions between Teleost Spinal Neurons: Electrophysiology and Ultrastructure , 1963, Science.

[5]  J T AITKEN,et al.  Neuron size and neuron population density in the lumbosacral region of the cat's spinal cord. , 1961, Journal of anatomy.

[6]  M. Ito,et al.  Electrical behaviour of the motoneurone membrane during intracellularly applied current steps. , 1965, The Journal of physiology.

[7]  J. Eccles,et al.  The relationship between the mode of operation and the dimensions of the junctional regions at synapses and motor end-organs , 1958, Proceedings of the Royal Society of London. Series B - Biological Sciences.

[8]  E. Gray,et al.  Comparative electron microscopy of synapses in the vertebrate spinal cord. , 1966, Journal of cell science.

[9]  P. G. Nelson,et al.  Anomalous rectification in cat spinal motoneurons and effect of polarizing currents on excitatory postsynaptic potential. , 1967, Journal of neurophysiology.

[10]  J. Eccles Membrane time constants of cat motoneurons and time courses of synptic action. , 1961, Experimental neurology.

[11]  D. Kernell,et al.  Input Resistance, Electrical Excitability, and Size of Ventral Horn Cells in Cat Spinal Cord , 1966, Science.

[12]  W. Rall Branching dendritic trees and motoneuron membrane resistivity. , 1959, Experimental neurology.

[13]  R. B. Wuerker,et al.  Membrane impedance changes during synaptic transmission in cat spinal motoneurons. , 1967, Journal of neurophysiology.

[14]  B. Katz,et al.  The measurement of synaptic delay, and the time course of acetylcholine release at the neuromuscular junction , 1965, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[15]  R. Schultz,et al.  Plasma Membrane Apposition in the Central Nervous System after Aldehyde Perfusion , 1964, Nature.

[16]  B. Katz,et al.  An analysis of the end‐plate potential recorded with an intra‐cellular electrode , 1951, The Journal of physiology.

[17]  R E Burke,et al.  Composite nature of the monosynaptic excitatory postsynaptic potential. , 1967, Journal of neurophysiology.

[18]  H. Mannen Contribution to the morphological study of dendritic arborization in the brain stem. , 1966, Progress in brain research.

[19]  S. Gelfan,et al.  Synaptic density on spinal neurons of normal dogs and dogs with experimental hind‐limb rigidity , 1964, The Journal of comparative neurology.

[20]  J. Eccles,et al.  Excitatory synaptic action in motoneurones , 1955, The Journal of physiology.

[21]  D. R. Curtis,et al.  The chemical excitation of spinal neurones by certain acidic amino acids , 1960, The Journal of physiology.

[22]  B. Katz,et al.  Propagation of electric activity in motor nerve terminals , 1965, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[23]  Wilfrid Rall,et al.  Theoretical significance of dendritic trees for neuronal input-output relations , 1964 .

[24]  W. Rall Membrane potential transients and membrane time constant of motoneurons. , 1960, Experimental neurology.

[25]  J. Eccles,et al.  The time courses of excitatory and inhibitory synaptic actions , 1959, The Journal of physiology.

[26]  W. Rall Distinguishing theoretical synaptic potentials computed for different soma-dendritic distributions of synaptic input. , 1967, Journal of neurophysiology.

[27]  W. Rall Electrophysiology of a dendritic neuron model. , 1962, Biophysical journal.

[28]  M. Kuno Quantal components of excitatory synaptic potentials in spinal motoneurones , 1964, The Journal of physiology.

[29]  Werner R. Loewenstein,et al.  STUDIES ON AN EPITHELIAL (GLAND) CELL JUNCTION I. Modifications of Surface Membrane Permeability , 1964 .