The avian hippocampus and the hypothetical maps used by navigating migratory birds (with some reflection on compasses and migratory restlessness)

The homology between the avian hippocampal formation (HF) and mammalian hippocampus nurtures the expectation that HF plays a fundamental role in navigation by migratory birds. Indeed, HF of migratory birds displays anatomical properties that differ from non-migratory species. Using a hypothetical framework of multiple maps of differing spatial resolution and range, homing pigeon data suggest that HF is important for navigating by landscape features near familiar breeding, over-wintering, and stop-over sites. By contrast, HF would be unimportant for an olfactory navigational map, which could be operational over unfamiliar space farther away from a goal location, nor is there any evidence for HF involvement in the sun or geomagnetic compass. The most intriguing question that remains open is what role HF may play in navigation when a migrant is thousands kms away from a familiar area, where homing pigeon data are uninformative and a geomagnetic map may be operational. Beyond navigation, successful migration depends on seasonal timing and often becoming nocturnally active. There is little evidence that HF plays a role in the timing of circannual and circadian cycles. Rather, circadian pacemakers including the pineal gland may control circadian timing of nocturnal restlessness and photoperiodic seasonal pacemakers likely control circannual expression.