Signal processing in a simple visual system: the locust ocellar system and its synapses.

The neurons with the widest axons that carry information into a locust brain belong to L-neurons, the large, second-order neurons of the ocelli. L-neurons play roles in flight control and boosting visual sensitivity. Their morphology is simple, and their axons convey graded potentials from the ocellus with little decrement to the brain, which makes them good subjects in which to study transmission of graded potentials. L-neurons are very sensitive to changes in light, due to an abnormally high gain in the sign inverting synapses they receive from photoreceptors. Adaptation ensures that L-neurons signal contrast in a light signal when average light intensity changes, and that their responses depend on the speed of change in light. Neurons L1-3 make excitatory output synapses with third-order neurons and with L4-5. These synapses transmit tonically, but are unable to convey hyperpolarising signals about large increases in light. Graded rebound spikes enhance depolarising responses. L1-3 also make reciprocal inhibitory synapses with each other and transmission at these decrements so rapidly that it normally requires a presynaptic spike. The resolution with which graded potentials can be transferred has been studied at the inhibitory synapses, and is limited by intrinsic variability in the mechanism that determines neurotransmitter release. Electron microscopy has shown that each excitatory connection made from an L-neuron to a postsynaptic partner consists of thousands of discrete synaptic contacts, in which individual dense-staining bars in the presynaptic neuron are associated with clouds of vesicles. Acetylcholine is likely to be a neurotransmitter released by L-neurons.