The development of physiological responses of the piriform cortex in rats to stimulation of the lateral olfactory tract.

Extracellular recording techniques in rats were used to follow the postnatal development of the evoked response of the piriform cortex to electrical stimulation of the lateral olfactory tract (LOT) from birth to adulthood. As in other species, LOT shock in adult rats produces short-latency activation of units in piriform cortex and an extracellular field potential consisting of three components: a surface-negative component, the A1 wave (corresponding to the cortical monosynaptic EPSP evoked by the LOT fibers); a second surface-negative component, the B1 wave (corresponding to reactivation of layer I dendrites by intracortical fibers); and a late surface-positive component, the period 2 wave. A conditioning shock 20-150 msec before the test shock profoundly inhibits both evoked unit activity and the B1 wave, while it facilitates the A1. At birth, units can be orthodromically activated by LOT stimulation in association with the A1 wave. There is also a surface-positive spikelike wave, the S wave, which represents the summation of cortical unit activity. The B1 wave is apparent early in the first postnatal week. However, in contrast to the prominent inhibition in the adults, for the first few days after birth, single-unit responses, multiple-unit activity, and the S wave are all facilitated by a preceding conditioning shock with intervals of 200 msec or less, in association with the facilitation of the A1 wave. A shift to inhibition is apparent with longer intershock intervals of 300-700 msec, which exceed the period during which paired shocks facilitate the A1 wave. During the remainder of the first two postnatal weeks,, partial suppression of evoked activity with intervals of less than 200 msec appears and progressively increases in strength, but inhibition at very long intershock intervals remains greater in magnitude. During this time, the duration of the inhibitory period also decreases to near the adult value of 200-300 msec. In the third postnatal week the pattern was similar to that in the adult, but the inhibition was still clearly weaker than in adults. These results suggest a delayed maturation of the cortical inhibitory circuitry; this conclusion has also been suggested by previously published observations in the developing neocortex and hippocampus. In addition, the acceleration with age of the conduction velocity of axons in the LOT was analyzed. The adult value of 9.6 m/sec was not achieved until some time after postnatal day 15, which parallels the myelinization of the tract as observed with the light microscope.