The shadow-induced withdrawal response, dermal photoreceptors, and their input to the higher-order interneuron RPeD11 in the pond snail Lymnaea stagnalis

The shadow-induced withdrawal response in Lymnaea stagnalis is mediated by dermal photoreceptors located on the foot, mantle cavity, and skin around the pneumostome area. Here, we determined whether we could obtain a neural correlate of the withdrawal response elicited by a shadow in a higher-order central neuron that mediates withdrawal behavior. We measured the electrophysiological properties of the higher-order interneuron Right Pedal Dorsal 11 (RPeD11), which has a major role in Lymnaea withdrawal behavior. In semi-intact preparations comprising the circumesophageal ganglia, the mantle cavity and the pneumostome, but not the foot and eyes, a light-on stimulus elicited a small short-lasting hyperpolarization and a light-off stimulus elicited a depolarization of RPeD11. We also determined that dermal photoreceptors make a monosynaptic contact with RPeD11. The dermal photoreceptor afferents course to the circumesophageal ganglia via the anal and genital nerves to the visceral ganglion, and/or via the right internal and external parietal nerves to the parietal ganglion. Finally, in addition to responding to photic stimuli, RPeD11 responds to both mechanical and chemical stimuli delivered to the pneumostome.