The anterior part of the piriform cortex (the APC) has been the focus of cortical-level studies of olfactory coding and associative processes and has attracted considerable attention as a result of a unique capacity to initiate generalized tonic-clonic seizures. Based on analysis of cytoarchitecture, connections, and immunocytochemical markers, a new subdivision of the APC and an associated deep nucleus are distinguished in the rat. As a result of its ventrorostral location in the APC, the new subdivision is termed the APC(VR). The deep nucleus is termed the pre-endopiriform nucleus (pEn) based on location and certain parallels to the endopiriform nucleus. The APC(VR) has unique features of interest for normal function: immunostaining suggests that it receives input from tufted cells in the olfactory bulb in addition to mitral cells, and it provides a heavy, rather selective projection from the piriform cortex to the ventrolateral orbital cortex (VLO), a prefrontal area where chemosensory, visual, and spatial information converges. The APC(VR) also has di- and tri-synaptic projections to the VLO via the pEn and the submedial thalamic nucleus. The pEn is of particular interest from a pathological standpoint because it corresponds in location to the physiologically defined "deep piriform cortex" ("area tempestas") from which convulsants initiate temporal lobe seizures, and blockade reduces ischemic damage to the hippocampus. Immunostaining revealed novel features of the pEn and APC(VR) that could alter excitability, including a near-absence of gamma-aminobutyric acid (GABA)ergic "cartridge" endings on axon initial segments, few cholecystokinin (CCK)-positive basket cells, and very low gamma-aminobutyric acid transporter-1 (GAT1)-like immunoreactivity. Normal functions of the APC(VR)-pEn may require a shaping of neuronal activity by inhibitory processes in a fashion that renders this region susceptible to pathological behavior.

"Surface-associated astrocytes" (SAAs) in posterior piriform cortex (PPC) are unique by virtue of a direct apposition to the cortical surface and large-caliber processes that descend into layer I. In this study additional unique and functionally relevant features of SAAs in PPC of the rat were identified by light and electron microscopy. Examination of sections cut parallel to the surface of PPC and stained for glial fibrillar acidic protein revealed that, in addition to descending processes, SAAs give rise to an extensive matrix of "superficial processes." Electron microscopy revealed that these superficial processes, together with cell bodies, form a continuous sheet at the surface of PPC with features in common with the glia limitans that is formed by endfeet in other cortical areas. These include a glia limiting membrane with basal lamina and similar associated organelles, including a striking array of mitochondria. Of particular interest, SAAs lack the domain organization observed in neocortex and hippocampus. Rather, superficial processes overlap extensively with gap junctions between their proximal regions as well as between cell bodies. Study of the descending processes revealed thin extensions, many of which appose synaptic profiles. We conclude that SAAs provide a potential substrate for bidirectional signaling and transport between brain and the pial arteries and cerebrospinal fluid in the subarachnoid space. We postulate that the spatially distributed character of SAAs in PPC reflects and supports the spatially distributed circuitry and sensory representation that are also unique features of this area.