Form, function and intracortical projections of spiny neurones in the striate visual cortex of the cat.

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Abstract
The neuronal circuitry and structure-function relationships of single neurons in the striate visual cortex of the cat were studied using a combination of electrophysiological and anatomical techniques. Glass micropipettes filled with horseradish peroxidase were used to record extracellularly from single neurons. After studying the receptive field properties, the afferent inputs of the neurons were studied by determining their latency of response to electrical stimulation at different positions along the optic pathway. Some cells were thus classified as receiving a mono- or polysynaptic input from afferents of the lateral geniculate nucleus (l.g.n.), via X- or Y-like retinal ganglion cells. Two striking correlations were found between dendritic morphology and receptive field type. All spiny stellate cells, and all star pyramidal cells in layer 4A, had receptive fields with spatially separate on and off subfields (S-type receptive fields). All the identified afferent input to these, the major cell types in layer 4, was monosynaptic from X- or Y-like afferents. Neurons receiving monosynaptic X- or Y-like input were not strictly segregated in layer 4 and the lower portion of layer 3. Nevertheless the X- and Y-like l.g.n. fibers did not converge on any of the single neuron so far studied. Monosynaptic input from the l.g.n. afferents was not restricted to cells lying within layers 4 and 6, the main termination zones of the l.g.n. afferents, but was also received by cells lying in layers 3 and 5. The projection pattern of cells receiving monosynaptic input differed widely, depending on the laminar location of the cell soma. This suggests the presence of a number of divergent paths within the striate cortex. Cells receiving indirect input from the l.g.n. afferents were located mainly within layers 2, 3 and 5. Most pyramidal cells in layer 3 had axons projecting out of the striate cortex, while many axons of the layer 5 pyramids did not. The layer 5 cells showed the most morphological variation of any layer, were the most difficult to activate by electrical stimulation, and contained some cells, which responded with the longest latencies of any cells in the striate cortex. This suggests that they were several synapses distant from the l.g.n. input. The majority of cells in layers 2, 3, 4 and 6 had the same basic S-type receptive field structure. Only layer 5 contained a majority of cells with spatially overlapping on and off subfields (C- and B-type receptive fields). The projection patterns of the cortical neurons do not necessitate a model of cortical organization where C- and B-type receptive fields are synthesized from S-type receptive fields. Most cells with axons leaving the striate cortex were no more than 1-3 synapses distant from the l.g.n. input. Within a small region of striate cortex there may be a number of circuits containing few neurons, perhaps with some common members, each processing some different aspect of the visual world.