Timeāplace learning in wild, free-living hummingbirds
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- print Zotero Link:: NA PDF:: NA Files:: ScienceDirect Snapshot Reading Note:: Maria C. Tello-Ramos, T. Andrew Hurly, Caitlin Higgott, Susan D. Healy 2015
Abstract
Animals can learn to revisit locations at which foraging resources will renew over time. This ability is known as ātimeāplaceā learning. Although there is clear evidence for timeāplace learning from animals trained and tested in the laboratory, not all species learn timeāplace tasks with similar ease. Since hummingbirds feed from food sources that are constant in space and that renew with time, it seems plausible these birds might learn timeāplace associations and do so readily.
Here, then, we tested whether wild, free-living rufous hummingbirds, Selasphorus rufus, could learn the time and place at which four artificial flower patches were rewarded. Flowers in each of the four patches contained reward for only 1h each day but the time and sequence in which patches were rewarded were repeated across days. Most birds learned when to visit each patch.
To determine whether the birds used ordinal or circadian timing to choose the correct patch at the correct time, we ended the experiment with a single test trial in which we presented the patches (all flowers empty) only at the fourth hour of the day. The birds visited neither the patch that was normally rewarded first (daily ordinal timing) nor the fourth patch (time of day). These results suggest that the time component of the timeāplace learning in these birds requires both ordinal and circadian information.
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Time-Place Learning
When an event is constant in space and time, animals should benefit from learning both the time and the place of that event. This ability is known as timeeplace learning. Typically in the laboratory a time-place learning protocol involves animals being trained to receive a food reward in one place at a specific time of day and a food reward in another place at another time.
To test whether wild, free-living hummingbirds would learn a timeeplace association, we presented territorial male rufous hummingbirds with four patches of artificial flowers whereby each of the patches was rewarded 1 h at a time (e.g. Patch 1 flowers contained a reward from 0800 to 0900 hours, Patch 2 flowers from 0900 to 1000 and so on for the four patches). Additionally, to determine which of two possible timing mechanisms birds might use to visit the correct flower patch at the correct time, after 6 days of training we gave the birds a test. During the test day we presented the patches only at the fourth hour (with all flowers empty). If birds learned the order in which they encountered rewarded patches they should go first to the flowers in the first patch of the day to contain a reward. If, however, the birds learned the specific times of day at which each patch contained a reward birds should fly directly to flowers in the fourth patch.
Trap Lines
Information that would allow them to avoid wasting energy by visiting unprofitable flowers should, therefore, be especially useful. Furthermore, for those hummingbirds that forage by following repeatable sequences around the same flowers (trap lines), timeeplace learning would seem especially advantageous.
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