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Journal of Neuroscience, Vol 15, 1617-1630, Copyright © 1995 by Society for Neuroscience
Learning and memory in the honeybee
M Hammer and R Menzel
Institut fur Neurobiologie, FU-Berlin, Germany.
Insects are favorable subjects for neuroethological studies. Their nervous
systems are relatively small and contain many individually identifiable
cells. The CNS is highly compartmentalized with clear separations between
multisensory higher order neuropiles in the brain and neuropiles serving
sensory-motor routines in the ventral cord (Huber, 1974). The rich behavior
of insects includes orientation in space and time, visual, chemical, and
mechanical communication, and complex motor routines for flying, walking,
swimming, nest building, defense, and attack. Learning and memory, though,
are not usually considered to be a strong point of insects. Rather, insect
behavior is often regarded as highly stereotyped and under tight control of
genetically programmed neural circuits. This view, however, does not do
justice to the insect order of Hymenoptera (bees, wasps, ants). Most
Hymenopteran species care for their brood either as individual females or
as a social group of females. Consequently, they regularly return to their
nest site to feed, protect, and nurse the larvae, store food, and hide from
adverse environmental conditions. Since they search for food (prey; nectar
and pollen on flowers) at unpredictable sites, they have to learn the
celestial and terrestrial cues that guide their foraging trips over long
distances and allow them to find their nest sites (central place foraging;
von Frisch, 1967; Seeley, 1985). They learn to relate the sun's position
and sky pattern of polarized light to the time of the day (Lindauer, 1959),
and landmarks are learned in relationship to the nest site within the
framework of the time- compensated sun compass. The honeybee communicates
direction and distance of a feeding place to hive mates by performing a
ritualized body movement, the waggle dance (von Frisch, 1967). Associative
learning is an essential component of the bee's central place foraging
behavior and dance communication. Hive mates attending a dance performance
learn the odor emanating from the dancing bee and seek it at the indicated
food site. The odor, color, and shape of flowers are learned when the bee
experiences these stimuli shortly before it finds food (nectar, pollen).
This appetitive learning in bees has many characteristics of associative
learning well known from mammalian learning studies (Menzel, 1985, 1990;
Bitterman, 1988). It follows the rules of classical and operant
conditioning, respectively, so that stimuli or behavioral acts are
associated with evaluating stimuli. Since associative learning, especially
of the classical type, is well described at the phenomenological and
operational level (Rescorla, 1988), it provides a favorable approach in the
search for the neural substrate underlying learning and memory.(ABSTRACT
TRUNCATED AT 400 WORDS)
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