Identification of a female sex pheromone in Carcinus maenas

Despite major progress in our understanding of animal signaling systems, the identification of chemical signals in aquatic organisms is still in its infancy. Exemplary for this is the lack of structural knowledge of crustacean sex pheromones despite their initial description in crabs almost 40 yr ago. We hypothesized that species in which the reproductive event is linked to a short window of mating opportunities centered around the time of ecdysis would be ideal to purify and identify such a cue. Here we present the first identification of a crustacean sex pheromone, the female signal produced by the green crab Carcinus maenas from both conditioned seawater and female urine. We used a bioassay-driven purification scheme combined with a candidate approach focusing on excreted compounds that changed through the course of the female molt cycle. Uridine diphosphate (UDP), a nucleotide accumulated from UDP-N-acetylglucosamine during chitin biosynthesis, is the major component of the female crab pheromone and induced all key characteristics of male sexual behavior in bioassays (i.e. pre- and post-copulatory guarding of the female and initiation of mating) at a threshold of 10(-5) M UDP in seawater. The identification of a nucleotide pheromone in crustaceans will enable future studies on signal specificity, biological function, biosynthesis, reception, and evolution as well as focus its potential use in pest control and aquaculture, thus allowing for a major expansion of crustacean chemical and behavioral ecology research that was hindered by the lack of structural knowledge.