f Wolbachia bacteria are endosymbionts that infect approximately 40% of all insect species and are best known for their ability to manipulate host reproductive systems. Though the effect Wolbachia infection has on somatic tissues is less well understood, when present in cells of the adult Drosophila melanogaster brain, Wolbachia exerts an influence over behaviors related to olfaction. Here, we show that a strain of Wolbachia influences male aggression in flies, which is critically important in mate competition. A specific strain of Wolbachia was observed to reduce the initiation of aggressive encounters in Drosophila males compared to the behavior of their uninfected controls. To determine how Wolbachia was able to alter aggressive behavior, we investigated the role of octopamine, a neurotransmitter known to influence male aggressive behavior in many insect species. Transcriptional analysis of the octopamine biosynthesis pathway revealed that two essential genes, the tyrosine decarboxylase and tyramine -hydroxylase genes, were significantly downregulated in Wolbachia-infected flies. Quantitative chemical analysis also showed that total octopamine levels were significantly reduced in the adult heads.
Male aggression is an important social behavior common throughout the animal kingdom. Aggressive behaviors are utilized to secure food, mates, and territory; these, in turn, ensure reproductive success and species propagation (1). An interplay between environmental influences, together with epigenetic and genetic factors (acting through neurotransmitters, hormones, and pheromones), shapes and influences behaviors in animals (2-6). Recently, evidence of the importance of microbial symbionts in directly manipulating host behavior, often by altering neurotransmitter levels in the brain (7-11), has changed the manner in which behavior is viewed. Inherited microbes exert such a strong effect that they, in addition to genetics/epigenetics and the surrounding environment, have been described as a third major determinant of behaviors (12).Wolbachia bacteria are reproductive endosymbiotic alphaproteobacteria and are the most common intracellular bacteria on Earth, infecting filarial nematodes, arachnids, and at least 40% of all insect species, including the model insect, Drosophila melanogaster (13-16). Wolbachia bacteria are maternally transmitted and are best known for their ability to manipulate host sex determination or reproductive systems in order to promote germ line transmission (17, 18). Wolbachia bacteria have also been shown to influence host metabolic pathways (19-21), provide protection from pathogens (22, 23), influence host life span (24), and play a role in host speciation events (25).Wolbachia infections have been correlated to changes in behaviors, including male mating frequencies (11), mate discrimination (26), and responses to olfactory cues (27)(28)(29). Wolbachia can also induce host selective mate avoidance, i.e., causing the host to avoid mates harboring another, incompatible symbiont variant, which ...