Pigeons remember visual landmarks after one release and rely upon them more if they are anosmic

Forty years ago, Papi and colleagues discovered that anosmic pigeons cannot find their way home when released at unfamiliar locations. They explained this phenomenon by developing the olfactory navigation hypothesis: pigeons at the home loft learn the odours carried by the winds in association with wind direction; once at the release site, they determine the direction of displacement on the basis of the odours perceived locally and orient homeward. In addition to the old classical experiments, new GPS tracking data and observations on the activation of the olfactory system in displaced pigeons have provided further evidence for the specific role of olfactory cues in pigeon navigation. Although it is not known which odours the birds might rely on for navigation, it has been shown that volatile organic compounds in the atmosphere are distributed as fairly stable gradients to allow environmental odour-based navigation. The investigation of the potential role of olfactory cues for navigation in wild birds is still at an early stage; however, the evidence collected so far suggests that olfactory navigation might be a widespread mechanism in avian species.

Wide-ranging animals, such as birds, regularly traverse large areas of the landscape efficiently in the course of their local movement patterns, which raises fundamental questions about the cognitive mechanisms involved. By using precision global-positioning-system loggers, we show that homing pigeons (Columba livia) not only come to rely on highly stereotyped yet surprisingly inefficient routes within the local area but are attracted directly back to their individually preferred routes even when released from novel sites off-route. This precise route loyalty demonstrates a reliance on familiar landmarks throughout the flight, which was unexpected under current models of avian navigation. We discuss how visual landmarks may be encoded as waypoints within familiar route maps.

GPS analysis of flight trajectories of pigeons can reveal that topographic features influence their flight paths. Recording electrical brain activity that reflects attentional processing could indicate objects of interest that do not cause changes in the flight path. Therefore, we investigated whether crossing particular visual landmarks when homing from a familiar release site is associated with changes in EEG. Birds carried both data-loggers for recording GPS position and EEG during flight. First, we classified characteristic EEG frequencies of caged birds and found five main bands: A: 0-3, B: 3-12, C: 12-60, D: 60-130, and E: 130-200 Hz. We analyzed changes in these activity bands when pigeons were released over sea (a featureless environment) and over land. Passing over the coastline and other prominent landmarks produced a pattern of EEG alterations consisting of two phases: activation of EEG in the high-frequency bands (D and/or E), followed by activation of C. Overlaying the EEG activity with GPS tracks allowed us to identify topographical features of interest for the pigeons that were not recognizable by distinct changes of their flight path. We provide evidence that EEG analysis can identify landmarks and objects of interest during homing. Middle-frequency activity (C) reflects visual perception of prominent landmarks, whereas activation of higher frequencies (D and E) is linked with information processing at a higher level. Activation of E bands is likely to reflect an initial process of orientation and is not necessarily linked with processing of visual information.