Primer pheromones play key roles in regulating division of labor, which is one of the most fundamental and defining aspects of insect sociality. Primer pheromones are chemical messengers that transmit hormone-like messages among colony members; in recipients these messages can either induce or suppress phenotypic caste differentiation. Here, we investigated soldier-caste-derived chemicals as possible primer pheromones in the lower termite Reticulitermes flavipes, a species for which no primer pheromones have yet been identified. We determined that soldier head extracts (SHE), when provided to totipotent workers along with the insect morphogenetic juvenile hormone (JH), significantly enhanced soldier caste differentiation. When applied alone, however, SHE had no impacts on caste differentiation, survivorship, or any other aspect of worker biology. These findings support that soldier-derived chemicals serve as primer pheromones which enhance the action of the endogenous morphogenetic hormone JH. Thus, SHE chemicals apparently have no effect when received under natural conditions by non-receptive individuals with presumably low JH titers. Gas chromatography-mass spectrometry analysis identified two terpenes as the most plentiful components of R. flavipes SHE. Through GC-MS and NMR analyses, these terpenes were identified as γ-cadinene and its corresponding aldehyde, g-cadinenal. Validative bioassays with commercially available cadinene confirmed its activity. However, several other previously identified terpenes were also significantly active. These findings reveal a novel primer pheromone-like function for soldier-derived terpenes in termites, and further suggest convergent evolution of terpene functions in enhancing JH-dependent soldier caste differentiation.


Termite; soldier; primer pheromone; juvenile hormone; terpene

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