Individual Learning
Much of an animal’s individual learning is via an associative mode; that is, an association is made between a stimulus and an outcome. Associative learning may arise either from classical (Pavlovian) conditioning, where an animal associates a biologically relevant stimulus (e.g., food) with a previously irrelevant stimulus (e.g., railway tracks), or from operant (instrumental) conditioning, where the behavior of the animal is controlled by the consequences of that behavior (e.g., feeding on grain on tracks leads to a food reward).
These learning processes can make a behavior more likely through positive reinforcement (via rewards) or negative reinforcement (via unpleasant stimuli), or less likely through punishment or inhibitory learning (again, via unpleasant stimuli). For example, a bear foraging on railway tracks (Murrayet al. 2017) might be more likely to forage when it receives grain rewards (positive reinforcement) but less likely to forage through negative interactions with moving trains (punishment or inhibitory learning). Additionally, it might increase its level of vigilance through negative interactions with moving trains (negative reinforcement).
One associative learning mode relevant to animal movement is discrimination learning, where an animal learns to respond differently to distinct stimuli. For example, because homing pigeons can discriminate between the presence and absence of anomalies in magnetic fields, magnetoreception could be used for navigation (Moraet al. 2004).
Two nonassociative learning modes that are relevant to movement are habituation (decreased response to a stimulus after repeated exposure) and sensitization (increased response to a stimulus after repeated exposure). These modes are related to the strength of association between stimulus and outcome, rather than the association itself. For example, the sensory responsiveness of honey bees declines after bees are subjected to low sucrose sugar solutions (habituation) and increases after bees are subjected to high sugar solutions (sensitization) (Scheiner 2004). In turn, the sensory responsiveness of honey bees constrains individual foraging plasticity and skews the collective foraging decisions of colonies (Scheiner 2004). Box 1 provides further details on these modes of learning.
Another mode of learning, latent learning, is also relevant to animal movement (Frankset al. 2007). Latent learning involves the gathering and storing of information, without immediate reward, such as when animals learn their migration route away from breeding grounds after they are born (e.g., in autumn) and must use that information to migrate back to breeding grounds in the spring.