Year of Publication


Degree Name

Doctor of Philosophy (PhD)

Document Type

Doctoral Dissertation


Agriculture, Food and Environment



First Advisor

Dr. Xuguo Zhou

Second Advisor

Dr. John J. Obrycki


The evolution of nonreproductive castes is a defining characteristic of eusociality. The function and developmental regulation of the altruistic worker and soldier caste is the central element contributing to major advantages of eusociality over solitary animals. The soldier caste is the first evolved sterile caste in termites. Their primary function is believed to be colony defense. However, the function and development of termite soldiers remains largely unknown. Because of their apparent morphological adaptation for fighting and their limited behavior repertoire, our understanding of colony defense by termite soldiers is limited to their physical defense. In addition, we know little about the molecular mechanisms mediating soldier development. In Chapters 2 and 3 I discuss the role of the soldier caste under competition risk. By exposing the Eastern subterranean termite Reticulitermes flavipes to cues of a competitor termite species, I found that exposure to competitor cues reduced feeding, compromised growth and survival of R. flavipes workers. The presence of R. flavipes soldiers largely ameliorated these negative impacts. At the transcriptional level, R. flavipes soldiers can counteract the effects of competitor cues on worker head gene expression. This counteracting effect seems to be associated with genes in metabolism and immunity. These studies demonstrate that competition can affect a termite colony’s fitness by either competitors physically invading the colony and causing damage or cues from competitors inducing a stress response in termite colony members. More importantly, soldiers can contribute to colony fitness by physically engaging in combat, but also by enhancing colony members’ survival under competitor-cue exposure. In Chapter 4, I describe the molecular mechanism mediating soldier-caste differentiation. I cloned the full length cDNA sequence of the R. flavipes Methoprene-tolerance (Met) gene, a gene encoding a putative receptor for juvenile hormones. Using RNA interference, I studied the function of Met and found that this gene essentially mediates the JH-dependent soldier-caste differentiation in termites.