Innervation patterns in rat saccular macula. A structural basis for complex sensory processing.

Serial sections through the anterior part of rat saccular macula were reconstructed as montages. Findings are that type II hair cells are integrated into the neural circuitry of type I cells, chiefly by synapses with neighboring calyces and their collaterals; and that complex interactions between afferent- and efferent-type nerve elements take place. Three basic types of nerve/calyx pattern are present: U-type nerves lose their myelin before they enter the macula and have complex calyces with several collaterals; M-type nerves are myelinated up to the calyx, which lacks collaterals; and M/U-type nerves have short, unmyelinated segments proximal to their calyces, which have few collaterals. Both afferent- and efferent-type collaterals spring from calyces, chiefly from those of U-type nerves. Type II cells are presynaptic both to electron-lucent and to vesiculated terminals; some synapses are reciprocal. Electron-lucent boutons sometimes are presynaptic to calyces and to type II hair cells; and morphologically afferent-to-afferent kinds of synapses occur in the neuroepithelium. The anatomical findings indicate that complex information processing must occur in mammalian gravity receptors.