The formation and maturation of synapses in the visual cortex of the rat. II. Quantitative analysis

We have examined quantitatively the formation and maturation of synapses in the visual cortex of the rat. The density of the total number of synapses (synapses per 100 micron2 neuropil) as well as the densities of Gray's type I and type II contacts were estimated from photographic montages of coronal strips of visual cortex from rats of various postnatal ages. Histograms of synaptic density as a function of depth were prepared, and the mean values of the postsynaptic density length and vesicle number per terminal were estimated for the two synapse types at each age examined. During the first few days of life, synapses were concentrated in the subplate region. By the latter part of the second postnatal week they were present throughout the cortex and an adult-like distribution, in which the highest densities were present in the superficial layers, was achieved by day 14. The postsynaptic density length of the type I synapses remained relatively unchanged during development but that of the type II synapses was more variable. Specifically, it was significantly longer during the second and third postnatal weeks compared to earlier ages and to adult values. The mean number of vesicles per terminal for the two synapse types increased with age until day 28. Subsequently, it only increased slightly between days 28 and 90 for the type I synapses but decreased significantly for the type II synaptic contacts. At all ages examined, type I synapses formed the majority of synaptic contacts. The developmental pattern appeared to differ for the two synapse types. The density of type I synapses increased continuously during the first three weeks and achieved a mean value close to that of adult animals by day 20. In contrast, the density of type II synapses did not increase significantly until day 6, increased dramatically in the second and third postnatal weeks, and then declined markedly between days 20 and 90. The observed decrease in the density of type II synaptic contacts is a clear example of synapse elimination in the visual cortex.