No abstract

The number of neurons, glia, and synapses in the visual cortex of adult macaque monkeys has been estimated by stereological methods. The data is presented separately for 13 sublaminae. For the total cortical thickness, the average numerical density of neurons is approximately 120,000 per mm3 of tissue. This density increases and decreases in the different sublaminae in the direction that one would expect from the classical descriptions of Nissl-stained material (e.g., 137,000/mm3 in IVC alpha; 211,000/mm3 in IVC beta). There are about 200,000 neurons under 1 mm2 or cortical surface: 28% are in layers I-III, 45% are in layer IV, and 27% are in layers V and VI. The total number in area 17 of one hemisphere is close to 160,000,000. For the total cortical thickness the numerical density of synapses is 276,000,000 per mm3 of tissue. Laminae with higher neuronal densities tend to have higher synaptic densities but the correlation is not perfectly concordant. Moreover, the changes in synaptic densities are not as great as those in neuronal densities so that laminae with higher neuronal densities have a lower synapse to neuron ratio and laminae with lower neuronal densities have a higher synapse to neuron ratio. For the total cortical thickness this ratio is 2,300 synapses per neuron (1,400 in layer IVC beta and 2,800 in layers II and III). There are 478 million synapses under 1 mm2 cortical surface: 40% are in layers I-III, 35% are in layer IV, and 25% are in layers V and VI. The numerical densities of astrocytes, oligodendrocytes, and microcytes are 38,000, 17,000, and 4,000 per mm3, respectively. The overall glia/neuron ratio is 0.49.

The number of synapses per unit volume of tissue (NV) has been estimated in individual laminae of the binocular and monocular regions of area 17 in six adult cats by using a method of size-frequency distribution. Separate estimates were obtained for RA synapses (containing round vesicles associated with asymmetric membrane differentiations) and for FS synapses (containing flat vesicles associated with symmetric membrane differentiations). For the total cortical thickness, the NV of all synapses is not statistically different between binocular (286 million per mm1(3] and monocular (281 million) regions, nor is it different between the two regions for any of the laminae. Eighty-four percent of synapses are of the RA type. Of those, 79% are found on dendritic spines, 21% on dendritic trunks, 0.1% on somata. FS synapses represent 16% of the total, with 31% of them on spines, 62% on dendritic trunks, and 7% on somata. The ratio of RA to FS synapses is kept relatively constant throughout the layers. A two-way analysis of variance shows no difference in the NV of either RA or FS synapses in the two regions nor in the NV or RA synapses between cats. It does, however, clearly demonstrate (p less than 0.001) interindividual differences for FS synapses. These variations between individual cats may be due to differences in age, breed, or environmental factors. In contrast to the relative uniformity of the NV of synapses between regions, the number of each type under 1 mm2 of cortical surface is 33% higher in the binocular region. This is due mainly to the greater thickness of the binocular region.

In order to study the mechanisms of synaptogenesis in the rat cerebellar cortex, a library of monoclonal antibodies has been generated against proteins of the isolated synapse. One recognizes a glycosylated 38 kDa protein that is concentrated in the synaptic vesicle fraction and resembles synaptophysin biochemically in its molecular weight, charge, and pattern of glycosylation. In the adult cerebellar cortex, the antisynaptophysin(mabQ155) immunoreactivity is codistributed with synapses. Immunoreactivity is strongest in the molecular layer where punctate deposits of reaction product outline the Purkinje cell dendrites. Discrete small profiles, consistent with the distribution of basket cell axon terminals, surround the Purkinje cells, and in the granular layer the synaptic glomeruli are intensely stained. There is no immunoreactivity in the white matter axon tracts. Electron microscope immunocytochemistry confirms the synaptic location of the antigen and suggests that the reaction product is associated with synaptic vesicles. Both round and flat vesicle populations are immunoreactive. Antisynaptophysin(mabQ155) has been used to follow synaptogenesis in the developing rat cerebellum. In the newborn rat (P0), despite the paucity of synapses, there is some specific immunoreactivity, especially in the subcortical white matter. Electron microscopy shows that the antigenicity is associated with vesicles within growth cones, filopodia, and immature axon profiles. During development, antisynaptophysin immunoreactivity increases progressively, along with the maturing cell populations, for both the granule cell-Purkinje cell and the mossy fiber-granule cell synapses. Quantitative biochemical analysis confirms the cytochemical results. These data suggest that neuronal growth cones express a synapse-specific antigen before complete morphological synapses are present.

The number of neurons in individual laminae of area 17 was determined separately for both the binocular and the monocular regions in the left hemisphere of six cats. The number of neurons/mm3 of tissue was obtained for each lamina by using the method of size-frequency distribution applied to neuronal nuclei. The number of neurons per unit of cortical surface could then be calculated from measurements of layer thickness. The number of neurons/mm3 of tissue for the total cortical thickness is on the order of 48,000 to 50,000 neurons, with no statistically significant differences between binocular and monocular regions. There are no significant differences for any of the layers except layer IV, in which the numerical density is 20% higher in the monocular region. The thickness of the cortex and of many of its layers, however, do vary between the two regions. Consequently there are significant differences in the number of neurons under 1 mm2 of surface. For the total cortical thickness there are significantly more (27%) neurons in the binocular (78,440) than in the monocular region (61,900). This overall difference is due to significant changes in layers 11, IIIA, IVA, and especially in layers V and VIA where neurons are 40% more numerous in the binocular region. These findings could signify either that the binocular region contains additional interneurons specifically related to binocular interactions or that it has a greater number of neurons projecting to other cortical and subcortical areas, or both.