Coding and adaptation during mechanical stimulation in the leech nervous system.

The experiments described here were designed to characterise sensory coding and adaptation during mechanical stimulation in the leech (Hirudo medicinalis). A chain of three ganglia and a segment of the body wall connected to the central ganglion were used. Eight extracellular suction pipettes and one or two intracellular electrodes were used to record action potentials from all mechanosensory neurones of the three ganglia. When the skin of the body wall was briefly touched with a filament exerting a force of about 2 mN, touch (T) cells in the central ganglion, but also those in adjacent ganglia (i.e. anterior and posterior), fired one or two action potentials. However, the threshold for action potential initiation was lower for T cells in the central ganglion than for those in adjacent ganglia. The timing of the first evoked action potential in a T cell was very reproducible with a jitter often lower than 100 us. Action potentials in T cells were not significantly correlated. When the force exerted by the filament was increased above 20 mN, pressure (P) cells in the central and neighbouring ganglia fired action potentials. Action potentials in P cells usually followed those evoked in T cells with a delay of about 20 ms and had a larger jitter of 0.5-10 ms. With stronger stimulations exceeding 50 mN, noxious (N) cells also fired action potentials. With such stimulations the majority of mechanosensory neurones in the three ganglia fired action potentials. The spatial properties of the whole receptive field of the mechanosensory neurones were explored by touching different parts of the skin. When the mechanical stimulation was applied for a longer time, i.e. 1 s, only P cells in the central ganglion continued to fire action potentials. P cells in neighbouring ganglia fully adapted after firing two or three action potentials.P cells in adjacent ganglia, having fully adapted to a steady mechanical stimulation of one part of the skin, fired action potentials following stimulation of a different region of the skin. These results indicate that a brief and localised stimulation of the skin can activate more than a dozen different mechanosensory neurones in the three ganglia and after 100 ms of steady stimulation many of these mechanosensory neurones stop firing action potentials and fully adapt. Adaptation occurs primarily at the nerve endings and mechanosensory neurones can quickly respond to mechanical stimulation at a different location on the skin.