Secondary olfactory projections and pallial topography in the Pacific hagfish, Eptatretus stouti.

The extent of the secondary olfactory projections shows great variation among different groups of craniates. Gnathostomes typically display restricted secondary olfactory projections, whereas lampreys have more extensive projections. Any attempt to determine the phylogenetic polarity of these characters, that is, to decide which is primitive and which is derived, requires an investigation of the secondary olfactory system in the sister group of lampreys and gnathostomes, the hagfishes. Therefore the secondary olfactory projections of the Pacific hagfish, Eptatretus stouti, were traced with the use of horseradish peroxidase and the lipophilic fluorescent tracing compound DiI. The projections are bilateral and massive to all pallial areas and the septum, moderate to the striatum, and relatively weak to the preoptic and infundibular regions of the hypothalamus, reaching caudally to the diencephalic-mesencephalic boundary. Afferents to the olfactory bulb arise from the pallium, the preoptic area, and the ventral thalamus. We compare the secondary olfactory projections in hagfishes with those in lampreys and in gnathostomes, and we conclude that the presence of extensive secondary olfactory projections is a primitive character of craniate brains.