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A long-lasting hyperpolarization has been observed in sensory neurones of the C.N.S. of the leech following natural stimulation applied to their receptive fields. The underlying mechanism of this hyperpolarization and its effects on signalling were investigated by intracellular recording.

1. The amplitude and duration of the hyperpolarization depended on the number of action potentials. Whereas a brief burst of impulses had little effect, a prolonged sensory discharge was followed by a hyperpolarization of up to 30 mV that gradually declined over a period of several minutes.

2. The hyperpolarization was abolished by ouabain, reversibly inhibited by strophanthidin or cooling, unaffected by Mg and accompanied by an increase in membrane resistance. These observations suggest that the hyperpolarization is the result of membrane current generated by an electrogenic pump.

3. A hyperpolarization similar to that recorded in the cell body also occurred in the neuronal processes within the neuropile, where synaptic contacts are made. This led to significant changes in integrative activity, such as an increase in the amplitude of excitatory synaptic potentials, a reversal of inhibitory synaptic potentials or a conduction block in parts of the neurones. All of these effects could be mimicked by injecting hyperpolarizing currents into resting cells, or abolished by injecting a depolarizing current into a cell hyperpolarized by previous impulses.

4. During neuronal hyperpolarization the sensitivity of the membrane potential to small increments of external K was enhanced by a factor of about three. This effect varied with the magnitude of the hyperpolarization produced by preceding impulses and could not be mimicked by applying K to a neurone hyperpolarized by the injection of current into its cell body.

5. These findings are discussed in relation to possible effects that changes in the intracellular and extracellular concentrations of ions produced by activity might have on integration in the C.N.S.

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