Properties of cholinergic responses in neurons in the intermediate grey layer of rat superior colliculus

The intermediate grey layer (SGI) of superior colliculus (SC) receives cholinergic innervation from brainstem parabrachial region. To clarify the action of cholinergic inputs to local circuits in the SGI, we investigated the effect of cholinergic agonists and antagonists on a large number of randomly sampled neurons in Wistar rat SGI (n=246) using whole-cell patch clamp technique in slices of the rat SC. Responses of the recorded cells (n=98) to bath application of carbachol were classified into five patterns: (i) nicotinic inward only (n=14); (ii) nicotinic inward+muscarinic inward (n=26); (iii) nicotinic inward+muscarinic inward+muscarinic outward (n=39); (iv) nicotinic inward+muscarinic outward (n=13) and (v) muscarinic outward only (n=4). Among these, a majority of morphologically identified projection neurons exhibited either response pattern (ii) (9/28) or (iii) (15/28), which suggested that the primary action of cholinergic inputs on the SGI output is excitatory. Nicotinic receptor subtypes involved in the nicotinic current were examined by testing the effects of antagonists on the currents induced by bath application of 1,1-dimethyl-4-phenyl-piperazinium or transient pressure application of acetylcholine (ACh). Muscarinic receptor subtypes involved in the muscarinic inward and outward currents were investigated by examining the effects of antagonists on muscarine-induced currents. The results showed that nicotinic inward currents are mediated mainly by alpha4beta2 and partly by alpha7 nicotinic receptors and that muscarinic inward and outward currents are mediated by M3 (plus M1) and M2 muscarinic receptors, respectively.