A Golgi study of the periaqueductal gray matter in the cat. Neuronal types and their distribution

In Golgi material, the neurons of the periaqueductal gray matter (PAG) of the cat have been classified into five types, according to the following criteria: number of dendrites per cell, characteristics of secondary arborization, frequency of spines and axon caliber. Type 1 cells, which are multipolar and rich in spines are the most frequent, and are probably intranuclear neurons. Type 4 cells have a short axon which ends in the PAG, but they differ from Type 1 in that their dendritic ramification is of a different type and there are few spines. Type 2 and 3 neurons have a thick axon which runs outside the PAG, and dendrites rich in spines. Type 2 cells have more primary dendrites, while Type 3 neurons have dendrites which may spread outside the PAG. Type 5 cells have dendrites with few spines and no secondary ramification. Their thick and long axon projects outside the PAG. Type 2, 3 and 5 cells have been considered projective neurons. The various neuron types are present in every area of the PAG, although in the ventral region there is a predominance of Type 2 and 5 neurons, in the dorsal regions of Type 2 and 3 cells, and in lateral regions of Type 3 and 5 cells. Local intrinsic circuits have been observed in which both the interneurons and the projective, with early axonic collaterals, are involved. The prevalence of neurons to which an afferent role has been attributed (Type 2 and 3 cells) compared with efferent cells (Type 5), is in agreement with hodological studies which indicate that the PAG receives multiple and numerous afferents in comparison with the relatively scarce efferent fibers. These projections can be intensely and deeply elaborated and modulated by means of local intrinsic circuits.