GABAergic interneurons containing calbindin D28K or somatostatin are major targets of GABAergic basal forebrain afferents in the rat neocortex

The arborization pattern and postsynaptic targets of the GABAergic component of the basal forebrain projection to neo‐ and mesocortical areas have been studied by the combination of anterograde tracing and pre‐ and postembedding immunocytochemistry. Phaseolus vulgaris leucoagglutinin (PHAL) was iontophoretically delivered into the region of the diagonal band of Broca, with some spread of the tracer into the substantia innominata and ventral pallidum. A large number of anterogradely labelled varicose fibres were visualized in the cingulate and retrosplenial cortices, and a relatively sparse innervation was observed in frontal and occipital cortical areas. Most of the labelled axons were studded with large en passant varicosities (Type 1), whereas the others (Type 2) had smaller boutons often of the drumstick type. Type 1 axons were distributed in all layers of the mesocortex with slightly lower frequency in layers 1 and 4. In the neocortex, layer 4, and to a smaller extent upper layer 5 and layer 6 contained the largest number of labelled fibres, whereas only a few fibres were seen in the supragranular layers. Characteristic type 2 axons were very sparse but could be found in all layers.