Intrinsic connections in cat visual cortex: a combined anterograde and retrograde tracing study

Area 18 of cat visual cortex was examined for intrinsic axons following small, columnar injections of an anterograde tracer,Phaseolus vulgaris leucoagglutinin (PHA-L). Locally projecting axons radiated from the injection site and branched to form 10–15 discrete, approximately circular patches 500–750 μm in diameter consisting of many bouton-studded terminal arborizations. Labeled fibers and boutons ramified densely in layers I, II/II, V, and VI, and were noticeably less dense in layer IV. Afferent and efferent pathways originating from the same cortical columns were studied by injecting a mixture of PHA-L and wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). Between 10 and 15 patches of cells retrogradely labeled by WGA-HRP surrounded each injection site. Within a patch, labeled cells were found in all layers and included both pyramidal and non-pyramidal cells. The distribution of PHA-L labeling was similar to that obtained when PHA-L was injected alone. Most often, the labeled patches resulting from injections of such mixtures contained both anterograde and retrograde labeling. However, patches consisting of retrograde labeling alone and of anterograde labeling alone were also observed, indicating that the local connections linking neighboring cortical columns were not always reciprocal.