Horizontal Interactions in Cat Striate Cortex: II. A Current Source-Density Analysis

The current source-density (CSD) analysis was used to investigate the organization of tangential synaptic connections in primary visual cortex of normally reared (NR) kittens and of NR, binocularly deprived (BD) and dark-reared (DR) adult cats. Laminar profiles of field potentials, elicited by intracortical microstimulation were measured at various distances from the stimulating electrodes. To exclude contamination by axon collaterals of antidromically stimulated thalamo-cortical fibres, these were destroyed by injecting the cytotoxin N-methyl-D-aspartate (NMDA) into the lateral geniculate nucleus 13 - 27 days before recording. The CSD profiles revealed distinct layer-specific patterns of lateral spread of activity. Invariably, the most prominent, long-lasting and far reaching responses were recorded in supragranular layers. Responses in layer IV were brief and confined to the vicinity of the stimulation site. Responses in infragranular layers spread as far as those in supragranular layers, but were of smaller amplitude. Latency considerations, the results of double shock stimulation, and the effects of translaminar cuts, suggest that these responses were monosynaptic and mediated by intracortical pathways with a conduction velocity of 0.3-5 m/s. The spatial spread of these responses changed substantially with age but was not influenced by visual deprivation. In NR adults, supra- and infragranular responses were recordable up to 2.5 mm from the stimulation site and layer IV responses up to 1 mm from the stimulation site. In kittens, the former responses spread up to 5 mm and the latter up to 2 mm from the site of stimulation. The amplitude of the responses decreased with distance from the stimulation site. This decrease was not always monotonic suggesting inhomogeneities in the tangential projections. The laminar distribution of current sinks and sources indicates that the pathways mediating tangential interactions form excitatory synapses on apical dendrites of pyramidal cells. It is concluded that the spatial spread of tangential excitatory interactions decreases with age, but that neither the laminar pattern nor the age-dependent reduction in the strength of tangential interactions are influenced by visual deprivation.