Dissection of the chemical
cues underlying courtship learning
The conditioned stimulus
(CS) for courtship conditioning has been assumed for many years to be
a component of the female's cuticular hydrocarbon. To determine what
chemicals constitute the CS, we trained males to discriminate between
mature and immature females and suppress courtship specifically to the
type of female that was associated with unsuccessful courtship (Ejima
et al., 2005). This "trainer-specific" learning is mediated by hydrocarbon
olfactory cues because it can be reconstituted with an extract of pheromone
and a generic courtship object. GC-MS analysis of the immature and mature
virgin hydrocarbon profiles revealed a limited number of compounds that
were specific to each type of female. These compounds are candidates
to be CSs. The nature of the unconditioned stimulus (US) for trainer-specific
learning is not clear, but appears to be the failure of the male to
copulate. In cases where the male was allowed to copulate during training,
no learning occurred and subsequent courtship was not suppressed. In
many other paradigms, failure to obtain an expected reward has been
shown to be aversive.
Training with a mated female
produces a qualitatively different outcome compared to training with
virgins: males show a generalized suppression of courtship towards all
types of females. This generalization of learning can be reproduced
by presentation of mated female pheromonal extracts during training
with a virgin. To determine what the chemical US for mated female conditioning
was, we compared mature virgin and mated female pheromonal profiles
by GC-MS. Mated females differ from age-matched virgins in only one
compound, cis-vaccenyl acetate (cVA). This hydrocarbon is produced only
by males, and is transferred to females during copulation in the ejaculate.
The presence of cVA during training with a virgin female can produce
generalized suppression of courtship, indicating that it is sufficient
to act as a US.
The effects of cVA on behavior,
however, appear to be context-dependent. We have found that the ability
to produce generalized courtship suppression varies with the nature
of the stimulatory pheromones that cVA is paired with. In addition,
other researchers have found that, at feeding sites, cVA can act as
a aggregation pheromone (Bartelt et al., 1985; Xu et al., 2005). Thus
the context and probably the processing of this olfactory cue can influence
its behavioral meaning to the animal.
Additional behavior topics: