Starving honey bee (Apis mellifera) larvae signal pheromonally to worker bees

Cooperative brood care is diagnostic of animal societies. This is particularly true for the advanced social insects, and the honey bee is the best understood of the insect societies. A brood pheromone signaling the presence of larvae in a bee colony has been characterised and well studied, but here we explored whether honey bee larvae actively signal their food needs pheromonally to workers. We show that starving honey bee larvae signal to workers via increased production of the volatile pheromone E-beta-ocimene. Analysis of volatile pheromones produced by food-deprived and fed larvae with gas chromatography-mass spectrometry showed that starving larvae produced more E-beta-ocimene. Behavioural analyses showed that adding E-beta-ocimene to empty cells increased the number of worker visits to those cells, and similarly adding E-beta-ocimene to larvae increased worker visitation rate to the larvae. RNA-seq and qRT-PCR analysis identified 3 genes in the E-beta-ocimene biosynthetic pathway that were upregulated in larvae following 30 minutes of starvation, and these genes also upregulated in 2-day old larvae compared to 4-day old larvae (2-day old larvae produce the most E-beta-ocimene). This identifies a pheromonal mechanism by which brood can beg for food from workers to influence the allocation of resources within the colony.