Developmental studies of thalamocortical and commissural connections in the rat somatic sensory cortex

Abstract

Autoradiographic, axonal degeneration, and horseradish peroxidase fiber tracing methods were employed to investigate the organization, development and potential plasticity of the thalamocortical projection to the somatic sensory cortex of the rat. In the adult animal, thalamocortical terminals are concentrated primarily in layers I and IV and in the upper part of layer VI. Fibers terminating in layers IV and VI arise from a different thalamic region than those terminating in layer I. Discrete clusters of fibers and terminals 250–450 μm wide are distributed only to the parts of the SI cortex containing dense aggregates of layer IV granule cells and not to the intervening, less granular and commissurally connected zones. At birth, thalamocortical fibers have invaded the deep part of the developing SI cortex and are concentrated in the upper part of layer VI. Between the age of two and three days, an additional concentration of fibers appears in the part of the cortical plate which will become layer IV. Layer IV is clearly recognizable by three days of age and the dense granule cell aggregates appear in it no more than one day later. The ingrowth of commissural fibers (Wise and Jones, '76) lags behind that of thalamic fibers. The mature commissural fiber pattern is not established until the age of seven days. After removal of the developing thalamocortical system by thalamotomy in newborn rats, subsequent investigation of the commissural system in the adult showed that no commissural fibers or terminals had invaded either laminae or zones of the cortex deprived of thalamic input. Similarly, commissurotomy at birth was not followed by sprouting of thalamic fibers into zones or laminae deprived of commissural connections. The connectional specificity observed in these neocortical fiber systems contrasts markedly with the plasticity of connections reported in allocortical systems. Removal of thalamocortical afferents before they attain their definitive distribution does not radically effect the overall development of the dense granule cell aggregates in layer IV. Within the aggregates, however, subsidiary features such as the “barrels” fail to appear. This finding suggests that certain elements of cortical architecture such as the dense granule cell aggregates are independent of thalamic afferents while others, such as the barrels, result from the interaction of the developing thalamocortical fibers and/or terminals with maturing neurons.