Substance P containing neurons was visualized by immunocytochemistry in the monkey hippocampus, subicular complex, and entorhinal cortex. Immunoreactive neurons were found solely in the hilar region of the dentate gyrus, and in strata oriens and pyramidale of Ammon's horn. In the subicular complex, immunoreactive neurons were located in those layers which were close to the alveus, whereas in the entorhinal cortex most of the substance P-positive neurons appeared in the second and third layers above the lamina dissecans. The majority of substance P-containing neurons were large multipolar cells, but small bipolar and multipolar cells also occurred in Ammon's horn, subiculum and entorhinal cortex. Dendrites of immunoreactive cells were smooth and displayed a few small, faintly stained spines which were hard to identify in the light microscopic preparations, but were visible with electron microscopy. Substance P-positive dendrites were exclusively found in the hilar region and never observed in the upper two-thirds of the molecular layer of the dentate gyrus. Moreover, immunoreactive dendrites rarely penetrated the stratum lacunosum-moleculare of Ammon's horn. In the electron microscopic preparations, somal and dendritic features of substance P-positive neurons were similar to those observed for GABAergic local circuit neurons. Axons of the substance P-immunoreactive local circuit neurons were thin and richly arborized in the upper two-thirds of the molecular layer of the dentate gyrus, in the stratum lacunosum-moleculare of Ammon's horn as well as in the subpial layers of the subicular complex and entorhinal cortex. Their terminals formed exclusively symmetric synapses with dendrites and spines. However, substance P-immunoreactive boutons were not found to make symmetric, axosomatic synapses on the granule cells of the dentate gyrus and very few were present on the pyramidal neurons of Ammon's horn, subicular complex, and entorhinal cortex. Hippocampal neurons, which were immunoreactive for substance P, also contained the neuropeptide somatostatin. However, not all of the somatostatin-containing neurons were substance P-immunoreactive. Thus, substance P-positive neurons are a subpopulation of somatostatin immunoreactive, GABAergic neurons. In conclusion, substance P-immunoreactive neurons are ideally suited for feed-back dendritic inhibition which may control the effectiveness of the main excitatory cortical input to the granule cells of the dentate gyrus and pyramidal neurons of the Ammon's horn.
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