Projections of the densocellular part of the hyperpallium in the rostral Wulst of pigeons (Columba livia).

The Wulst in birds shows a four-layered structure: apical part of the hyperpallium (HA), interstitial part of HA (IHA), intercalated part of hyperpallium (HI), and densocellular part of hyperpallium (HD). The Wulst consists of a small rostral somatosensory region and a larger caudal visual region. The visual HD relays visual information to IHA and HA in the Wulst and also transfers visual information to the hippocampal formation (Atoji et al., J Comp Neurol 526: 146-165, 2018). However, fiber pathways of the rostral HD remain unknown. In the present study, the fiber connections of the rostral HD and overlying HI were analyzed with tract-tracing techniques using a combination of injections of cholera toxin subunit B (CTB) for retrograde tracing and biotinylated dextran amine (BDA) for anterograde tracing. When the two tracers were bilaterally but separately injected into the rostral HD, major reciprocal connections were seen with the rostral HA, prepiriform cortex, and subdivisions of the hippocampal formation. One-way projections of huge fibers also reached the medial part of the medial striatum. When CTB and BDA were bilaterally and separately injected into the rostral HI, strong reciprocal connections were found between the rostral HI and HA, and weak connections were seen with areas outside the Wulst. These results suggest that the fiber pathways of the rostral HD and HI are distinguishable from each other in the telencephalon and suggest also that the rostral HD relays information to the rostral HA and simultaneously acts as a mediator to the hippocampal formation.