Distance Estimation by Foraging Honeybees

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Abstract
Honeybees are widely believed to assess feeder distances by the energy spent on foraging flights. However, a critical review of this 'energy hypothesis' reveals many inconsistencies in the experiments from which it was derived. In fact, new evidence shows that the energy hypothesis cannot be correct. Foragers loaded with weights do not overestimate distance, as indicated by their waggle dances performed upon return to the hive. Bees that climb to a feeder on top of a high building (50 m) signal the same distance as hive mates that visit an equidistant feeder at ground level. Foragers visiting a feeder suspended from a balloon at 70 m from their hive underestimate the distance flown dramatically when the balloon lifts the feeder from ground level to 90 m, even though the energy required to reach the feeder increases considerably. Foragers from a hive situated on a high building (50 m) that fly to a feeder on the roof of another high building (34 m) signal a much shorter distance than the actual distance flown. We propose instead an 'optical flow hypothesis': bees use the speed of retinal image motion perceived from the ground to estimate the distance flown. Flight altitude is important for distance estimation by retinal image flow, because objects move faster and farther across the retina when the bee flies closer to the ground. When the forager's flight behavior is considered, the optical flow hypothesis does not conflict with previous findings.