S. H. C. Hendry

Morphological diversity of immunocytochemically identified GABA neurons in the monkey sensory-motor cortex

SummaryGABAergic neurons have been identified in monkey sensory-motor cerebral cortex by light microscopic, immunocytochemical localization of the GABA synthesizing enzyme, glutamic acid decarboxylase (GAD). All GAD-positive neurons are non-pyramidal cells. Their somata are present in all layers and are evenly distributed across layers II-VI of the motor cortex (area 4), but are found in greater concentrations in layers II, IV and VI of all areas of first somatic sensory cortex (SI; areas 3a, 3b and 1–2). GAD-positive puncta (putative axon terminals) are present throughout the sensory-motor cortex, and they are found immediately adjacent to the somata, dendrites and presumptive axon initial segments of GAD-negative pyramidal cells. In addition, they are observed in close approximation to the somata of both large and small GAD-positive neurons. In area 4, the density of puncta is highest in the superficial cortical layers (layers I-III) and gradually declines throughout the deeper layers. In SI, the highest densities of puncta are present in layer IV, while moderately high densities are found in layers I-III and VI. In areas 3a and 3b, the puncta in layers IV and VI are particularly numerous and form foci that exhibit greater density than adjacent regions.GAD-positive neurons withlarge somata, 15–33 μ in diameter, are present in layers IIIB-VI of all areas. Such cells have many primary dendrites that radiate in all directions. In addition they have axons that ascend either from the superficial aspect of the somata or from primary dendrites, and that exhibit horizontal collateral branches. These neurons closely resemble the large basket cells (Marin-Padilla, 1969; Jones, 1975), and they may give rise to many of the GAD-positive endings surrounding the somata and proximal dendrites of pyramidal cells in layers III-VI. In addition,small GAD-positive somata are present in all layers, but they are most numerous in layers II and IIIA of all areas and in layer IV of SI. The somata and proximal dendrites of these cells vary from a multipolar shape with small, beaded dendrites, found primarily in layer IV, to bitufted and multipolar shapes with larger, smooth dendrites. The diversity of somal sizes and locations, the variety of dendritic patterns, and the different distributions of GAD-positive puncta, all combine to suggest that several different morphological classes of intrinsic comprise the GABA neurons of monkey cerebral cortex.

Synaptic organization of immunocytochemically identified GABA neurons in the monkey sensory-motor cortex

SummaryNeurons in the monkey somatic sensory and motor cortex were labelled immunocytochemically for the GABA synthesizing enzyme, glutamic acid decarboxylase (GAD), and examined with the electron microscope. The somata and dendrites of many large GAD-positive neurons of layers III–VI receive numerous asymmetric synapses from unlabelled terminals and symmetric synapses from GAD-positive terminals. Comparisons with light and electron microscopic studies of Golgi-impregnated neurons suggest that the large labelled neurons are basket cells. Small GAD-positive neurons generally receive few synapses on their somata and dendrites, and probably conform to several morphological types. GAD-positive axons form symmetric synapses on many neuronal elements including the somata, dendrites and initial segments of pyramidal cells, and the somata and dendrites of non-pyramidal cells. Synapses between GAD-positive terminals and GAD-positive cell bodies and dendrites are common in all layers. Many GAD-positive terminals in layers III–VI arise from myelinated axons. Some of the axons form pericellular terminal nests on pyramidal cell somata and are interpreted as originating from basket cells while other GAD-positive myelinated axons synapse with the somata and dendrites of non-pyramidal cells. These findings suggest either that the sites of basket cell terminations are more heterogeneous than previously believed or that there are other classes of GAD-positive neurons with myelinated axons. Unmyelinated GAD-positive axons synapse with the initial segments of pyramidal cell axons or formen passant synapses with dendritic spines and small dendritic shafts and are interpreted as arising from the population of small GAD-positive neurons which appears to include several morphological types.

Thalamic inputs to identified commissural neurons in the monkey somatic sensory cortex

SummaryCommissurally projecting neurons were identified in the monkey first somatic sensory area (SI) by the retrograde axonal transport of horseradish peroxidase (HRP) injected into the contralateral cortex. Neurons identified in this way have large pyramidal somata primarily in layer IIIB of the SI area. Their basal dendrites lie within the terminal plexus of thalamocortical afferents.Electron microscopy was used to examine the synaptic relations of the labelled commissural cells, in particular to determine whether they receive monosynaptic thalamic connections. To do this, retrogradely labelled commissural cells and Golgi-impregnated large pyramidal neurons from layer IIIB were examined ultrastructurally in material in which thalamocortical terminals were degenerating due to a prior lesion of the thalamus. In a significant number of cases degenerating terminals were found to make synapses on the spines or shafts of labelled dendrites.Injections of HRP into SI or into the white matter adjacent to the corpus callosum labelled callosal axons and terminals in the opposite SI. These axons terminated mainly near the somata of the layer IIIB pyramidal cells. Some of their terminals were found to synapse with dendrites receiving synaptic contacts from thalamocortical axon terminals.