The development of lamination of afferent fibers to the olfactory cortex in rats, with additional observations in the adult.

The complementary distribution of the fibers from the olfactory bulb and the intracortical associational fibers to layers Ia and Ib, respectively, of the olfactory cortex has been examined in both adult and neonatal rats, using horseradish peroxidase (HRP) and 3H-leucine as double tracers in the same animal. The observations presented here confirm and extend the previous demonstration (Price, '73) that in the adult the two projections are essentially nonoverlapping throughout the olfactory cortex. Indeed, when the distribution of axons from the olfactory bulb (labeled by HRP inserted into a cut in the LOT) is compared on the same section with that of associational fibers (labeled by 3H-leucine injected into the cortex), the overlap between the two projections is limited to a zone only 5-10 micron in width in both the piriform cortex and olfactory tubercle. In contrast, at P1 the two projections overlap throughout layer I, although the bulbar and associational fibers are slightly concentrated superficially and deeply in layer I, respectively. This overlap is especially prominent in the part of the anterior piriform cortex deep to the LOT. During the remainder of the first postnatal week, this overlap resolves and by P7 the segregation of the two sets of afferent fibers is nearly equivalent to that seen in the adult. However, there are several instances in adults where the segregation of these afferents does not develop. First, a relatively small population of aberrant axons derived from the LOT may be traced from layer Ia into layer Ib and then back to layer Ia. Most of these axons are large in diameter and lack the boutonlike varicosities found on smaller axons in layer Ia. They are most prominent in areas where the cortex is highly curved. Second, in layer I of the nucleus of the lateral olfactory tract, bulbar and associational fibers are extensively intermingled. In this case also, the bulbar fibers are large in diameter with only a few boutonlike varicosities. The developmental emergence of afferent segregation and its breakdown in cases where the fibers from the olfactory bulb do not form boutons suggest that an interaction between the two distinct sets of fibers and the dendritic field is responsible for the normal development of this segregation and that this interaction depends on the process of synaptogenesis.