Layer VI in cat primary auditory cortex: Golgi study and sublaminar origins of projection neurons

The organization of layer VI in cat primary auditory cortex (AI) was studied in mature specimens. Golgi-impregnated neurons were classified on the basis of their dendritic and somatic form. Ipsilateral and contralateral projection neurons and the corticogeniculate cells of origin were labeled with retrograde tracers and their profiles were compared with the results from Golgi studies. Layer VI was divided into a superficial half (layer VIa) with many pyramidal neurons and a deeper part (layer VIb) that is dominated by horizontal cells. Nine types of neuron were identified; four classes had subvarieties. Classical pyramidal cells and star, fusiform, tangential, and inverted pyramidal cells occur. Nonpyramidal neurons were Martinotti, multipolar stellate, bipolar, and horizontal cells. This variety of neurons distinguished layer VI from other AI layers. Pyramidal neuron dendrites contributed to the vertical, modular organization in AI, although their apical processes did not project beyond layer IV. Their axons had vertical, intrinsic processes as well as corticofugal branches. Horizontal cell dendrites extended laterally up to 700 microm and could integrate thalamic input across wide expanses of the tonotopic domain. Connectional experiments confirmed the sublaminar arrangement seen in Nissl material. Commissural cells were concentrated in layer VIa, whereas corticocortical neurons were more numerous in layer VIb. Corticothalamic cells were distributed more equally. The cytological complexity and diverse connections of layer VI may relate to a possible role in cortical development. Layer VI contained most of the neuronal types found in other layers in AI, and these cells form many of the same intrinsic and corticofugal connections that neurons in other layers will assume in adulthood. Layer VI, thus, may play a fundamental ontogenetic role in the construction and early function of the cortex.