The visual claustrum of the cat. I. Structure and connections.

The cat's dorsocaudal claustrum was studied in Golgi preparations, by electron microscopy, and by anterograde and retrograde tracer techniques. It receives a convergent retinotopic projection from several visual cortical ares, including areas 17, 18, 19, 21a and PMLS (posteromedial lateral suprasylvian area). The projection arises from spiny dendrite cells (pyramidal and fusiform) in the middle of cortical layer VI. As shown by a double label experiment, they form a separate population from those projecting to the lateral geniculate nucleus. There are also inputs from the lateral hypothalamus, from the nucleus centralis thalami, and probably from the locus coeruleus, but not from the sensory nuclei of the thalamus. Non-visual cortical areas do not project to the visual claustrum, but many of them are connected to other parts of the nucleus. For example, the splenial (cingulate) gyrus projects to a claustral zone just ventral to the visual area, and regions anterior to the visual area are connected with somatosensory and auditory cortex. The commonest cell type in the claustrum is a large spiny dendrite neuron whose axon leaves the nucleus after giving off local collaterals. Small spine-free cells, with beaded dendrites and a locally arborizing axon, are found also. Electron microscopy of the claustrum after ablation of the visual cortex showed degenerating type 1 axon terminals synapsing on spines and beaded dendrites, suggesting a direct cortical input to both cell types. The visual claustrum projects back to the visual cortex, to the same areas from which it receives an input. The return projection is predominantly ipsilateral, but there is, in addition, a small crossed projection. The claustrocortical axons terminate in all cortical layers but most heavily in layers IV and VI. The majority of the cells in the visual claustrum project to the cortex, and retinotopy is maintained throughout the entire corticoclaustral loop. No subcortical projections from the claustrum could be identified.