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.