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The Journal of Neurophysiology Vol. 84 No. 1 July 2000, pp. 39-50
Copyright ©2000 by the American Physiological Society
Department of Neurosciences, Case Western Reserve University, Cleveland, Ohio 44106
Friedman, Daniel and
Ben W. Strowbridge.
Functional Role of NMDA Autoreceptors in Olfactory Mitral Cells. J. Neurophysiol. 84: 39-50, 2000. The output of the olfactory bulb is governed by the interaction
of synaptic potentials with the intrinsic conductances of mitral cells.
While mitral cells often are considered as simple relay neurons,
conveying activity in olfactory receptor cells to the piriform cortex,
there is strong physiological and behavioral evidence that local
synaptic interactions within the olfactory bulb modulate mitral cell
discharges and facilitate odorant discrimination. Understanding the
circuitry of the olfactory bulb is complicated by the fact that most
dendrites in this region are both pre- and postsynaptic. Feedback
inhibition is mediated through reciprocal dendrodendritic synapses
between the secondary dendrites of mitral cells and GABAergic granule
cells. Here we show that glutamate released from mitral cell dendrites
also activates local N-methyl-D-aspartate (NMDA)
autoreceptors, generating an inward tail current following depolarizing
voltage steps. Autoreceptor-mediated self-excitation is calcium
dependent, can be evoked by single action potentials in the presence of
magnesium, and is graded with the number of spikes in a train. We find
that dendrodendritic inhibition also is evoked by single action
potentials but saturates rapidly during repetitive discharges.
Self-excitation also underlies the prolonged afterdischarges apparent
in mitral cells following potassium channel blockade. Both
afterdischarges and autoreceptor-mediated tail currents persist in TTX,
suggesting that they are produced by local rather than polysynaptic
actions of glutamate. Blockade of NMDA autoreceptors with
2-amino-5-phosphonovaleric acid (APV) reduces the firing frequency
within action potential cluster. The rapid kinetics of self-excitation
suggests a functional role of NMDA autoreceptors in prolonging periods
of phasic firing in mitral cells.
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