Time-Place learning

Time-place learning (TPL) is the process by which animals link events (e.g. finding food, encountering a predator) with both the location and time of occurrence. It enables them to decide which locations to visit or to avoid based on previous experience and knowledge of the current time of day. TPL presumably allows animals to maximize their chances of finding resources (food, mates) and avoiding predators, increasing survival chances. TPL requires spatial memory and a sense of time. The latter may be based on external time-cues (Zeitgebers), or internally generated circadian rhythms ("biological clock"). TPL may fundamentally underlie episodic memory.

The first evidence for time-place learning in animals came from studies in the 1930s on honeybees, which could be trained to visit two different feeders, one in the morning and the other in the afternoon. Subsequent work in the 1980s showed that only a few individuals in the colony were able to learn that task, and did so with more precision for the morning than for the afternoon feeding. Honeybees can also be trained to recognize one visual pattern to obtain food in the morning, and another pattern to get food in the afternoon; when presented with both patterns simultaneously, the same bees choose the "morning" pattern in the morning and the "afternoon" pattern in the afternoon.

The neotropical ant Ectatomma ruidum is also capable of time-place learning. They can learn to visit a feeding site in the morning, another one at midday, and a third one in the afternoon. They follow this spatio-temporal pattern even when food is withheld on test days. They then stay at the feeder for the approximate duration that food was normally available before moving on, at the right time, to the next feeder. This shows that the ants do not rely on direct cues from the food itself, but use instead a learned association between daily time and place.

In fish, time-place learning has been demonstrated in the golden shiner and the inanga. Golden shiners could be taught to seek food in one half of their aquarium in the morning, in the other half at midday, and back to the first half in the afternoon. They maintained this spatio-temporal pattern even when food was withheld on test days. The spatio-temporal pattern also shifted gradually over several days when the day-night cycle was suddenly shifted early by 6 h, as is typical of circadian rhythms. Inangas could be taught to seek food in one half of their aquarium in the morning and in the other half in the afternoon, even on test days when food was withheld; however, they could not be taught to avoid one half in the morning and the other half in the afternoon in response to simulated attacks by a heron.

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