Juvenile hormone and hemimetabolan eusociality: a comparison of cockroaches with termites

Termites are social Dictyoptera that evolved eusociality independently from social Hymenoptera. They are characterized by unique developmental plasticity that is the basis of caste differentiation and social organization. As developmental plasticity is a result of endocrine regulation, in order to understand the evolution of termite sociality it is helpful to compare the endocrine underpinning of development between termites and cockroaches. Nijhout and Wheeler (1982) proposed that varying JH titers determine caste differentiation in termites. Based on current results, we extend this model by adding the importance of social interactions. High JH titers in the presence of soldiers lead to regressive development (decrease in body size, apparent regression in development), while an absence of soldiers induces (pre-)soldier differentiation. On the opposite side, low JH titers in colonies headed by reproductives result in progressive molts toward adults, while an absence of reproductives induces development of replacement reproductives. In cockroaches, transcription factors involved in JH signaling, including the adult specifier E93 (the co-called MEKRE93 pathway) regulate the morphogenetic transition between the nymph and the adult. In termites, we speculate that castes might be determined by social effects playing a modulatory action of JH in the MEKRE93 pathway.