Multisensory response modulation in the superficial layers of the superior colliculus.

The mammalian superior colliculus (SC) is made up of seven distinct layers. Based on overall differences in neuronal morphology, afferent and efferent projection patterns, physiological properties, and presumptive behavioral role, the upper three layers have been classically grouped together as the superficial layers and the remaining four layers collectively make up the deep layers. Although the superficial layers receive their primary inputs from the retina and primary visual cortex, the deep layers receive inputs from extrastriate visual cortical areas and from auditory, somatosensory, and motor-related structures. In contrast, there is no evidence of monosynaptic nonvisual inputs to the superficial layers. However, more recent studies have revealed anatomical connections between the superficial and deep layers, thus providing the substrate for possible communication between these two functional divisions of the SC. In this study, we provide physiological evidence for auditory influences on visual responses in the superficial layers of the SC. Using extracellular recordings of local field potentials (LFPs) and multiunit activity, we demonstrate multisensory effects in the superficial layers of the cat SC such that subthreshold auditory activity (as seen in the LFP) modulates visual responses (reflected in spiking activity) when the two stimuli are presented together. These results have important implications for our understanding of the functional organization of the SC and for the neural basis of multisensory integration in general.