Cholecystokinin-like immunoreactive neurons in rat cerebral cortex.
The distribution and form of cholecystokinin immunoreactive neurons in neocortical areas within the posterior pole of the rat cerebral hemisphere was examined using the immunoperoxidase technique. Although cholecystokinin-positive neurons are present throughout the cortex, they are most frequent in the supragranular layers. These neurons are of three kinds: layer I neurons, bipolar cells, and other non-pyramidal cells with either multipolar or bitufted dendritic trees. In electron-microscopic preparations, the horseradish peroxidase reaction product is found to form a granular deposit which occurs throughout the cytoplasm and nucleoplasm and shows no predilection for any particular type of organelle. Electron-microscopy also shows cholecystokinin-positive neurons to have both symmetric and asymmetric synapses on their perikarya, which is additional evidence in favor of the interpretation that they are non-pyramidal cells. In the light-microscopic preparations three types of CCK-positive axons are encountered. These are vertically-oriented axons considered to arise from bipolar cells, a plexus in the superficial portion of layer II/III which is believed to arise from the multipolar and bitufted cells, and a deep plexus of unknown origin in layers VI and V. Since the axons of bipolar cells form asymmetric synapses they are thought to be excitatory neurons. In contrast, the bitufted and multipolar neurons are probably inhibitory, for previous studies have shown neurons with similar features to have axons which form symmetric synapses and to contain glutamic acid decarboxylase. Thus, although iontophoretically-applied cholecystokinin excites cortical neurons, it appears to be present in some neurons which are excitatory and others which are inhibitory.