Neocortical gamma-aminobutyric acid (GABA)ergic neurons have been previously described as largely involved in local intracortical circuitry. However, our recent findings in the murine model described select neocortical GABAergic neurons that project to both neighboring and more distant neocortical regions. Here, we investigated whether such GABAergic projection neurons are also found in the cat neocortex. Wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP) was injected into the visual, auditory, or somatosensory cortex, in order to label efferent cortical neurons retrogradely and to label axons and terminals orthogradely. Staining for nicotinamide adenine dinucleotide phosphate diaphorase (NADPH-d), an enzyme involved in nitric oxide synthesis, was employed, and co-localization with WGA-HRP was determined by means of both polarizing and brightfield microscopy. We concluded that neurons double-labeled with WGA-HRP and NADPH-d in a distant region from the WGA-HRP-injection site are GABAergic neurons with long-range projection axons. All double-labeled neurons were found in cortical layers VIa and VIb and in the white matter. Neurons with intense NADPH-d reactivity (type I) were determined to be neuronal nitric oxide synthase (nNOS) positive in all cases. However, weakly NADPH-d-reactive neurons (type II) lacked nNOS immunoreactivity. Moreover, nNOS often co-localized with GABA, neuropeptide-Y, and somatostatin in the cat neocortex. In summary, the GABAergic neurons described here projected in a manner similar to that previously described for neocortical principal neurons, although some unique GABAergic long-range projections were also demonstrated.
γ-aminobutyric acid (GABA)ergic neurons in the neocortex have been regarded as interneurons and speculated to modulate the activity of neurons locally. Recently, however, several experiments revealed that neuronal nitric oxide synthase (nNOS)-positive GABAergic neurons project cortico-cortically with long axons. In this study, we illustrate Golgi-like images of the nNOS-positive GABAergic neurons using a nicotinamide adenine dinucleotide phosphate diaphorase (NADPH-d) reaction and follow the emanating axon branches in cat brain sections. These axon branches projected cortico-cortically with other non-labeled arcuate fibers, contra-laterally via the corpus callosum and anterior commissure. The labeled fibers were not limited to the neocortex but found also in the fimbria of the hippocampus. In order to have additional information on these GABAergic neuron projections, we investigated green fluorescent protein (GFP)-labeled GABAergic neurons in GAD67-Cre knock-in/GFP Cre-reporter mice. GFP-labeled axons emanate densely, especially in the fimbria, a small number in the anterior commissure, and very sparsely in the corpus callosum. These two different approaches confirm that not only nNOS-positive GABAergic neurons but also other subtypes of GABAergic neurons project long axons in the cerebral cortex and are in a position to be involved in information processing.
Ascending projections from the nucleus of the brachium of the inferior colliculus (NBIC) in the cat were studied by the autoradiographic tracing method. Many fibers from the NBIC ascend ipsilaterally in the lateral tegmentum along the medial border of the brachium of the inferior colliculus. At midbrain levels, fibers from the NBIC end in the superior colliculus, the pretectum, the central gray and the peripeduncular tegmental region bilaterally with ipsilateral predominance. NBIC fibers to the superior colliculus are distributed densely to laminae VI and III throughout the whole rostrocaudal extent of the colliculus. In the pretectum, NBIC fibers terminate in the anterior and medial nuclei and the nucleus of the posterior commissure. NBIC fibers to the dorsal thalamus are distributed largely ipsilaterally. Many NBIC fibers end in the dorsal and medial divisions of the medial geniculate body, but few in the ventral division. The NBIC also sends fibers to the suprageniculate, limitans and lateralis posterior nuclei and the lateral portion of the posterior nuclear complex; these regions of termination of NBIC fibers constitute, as a whole, a single NBIC recipient sector. Additionally, the NBIC sends fibers to the centralis lateralis, medialis dorsalis, paraventricular and subparafascicular nuclei of the thalamus.
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