Queen bee and her retinue. Photo by Kathy Keatley Garvey

DAVIS, CA (MPG) – Noted bee scientist Robert E. Page Jr. examines the nurse-to-forager transition in honey bees in an entirely different perspective in newly published research in the Proceedings of the National Academy of Sciences (PNAS).

Page, a retired UC Davis and Arizona State University (ASU) entomology professor and administrator, along with scientist Eric Bonabeau of Berkeley, analyzed “the nurse-to-forager transition in worker honey bees (Apis mellifera) from a network perspective, treating the transcriptome as a gene–gene correlation system whose global organization reflects collective regulatory states.” 

“Division of labor in social insects is enabled by coordinated changes in physiology, neural signaling and gene regulation, yet it remains unclear to what extent behavioral maturation reflects incremental changes in individual genes and/or a reorganization of transcriptomic regulation at the system level,” they explain in the paper, titled “System-Level Reorganization of Gene Networks Underlies the Nurse-to-Forager Transition in Honey Bees.”

Page wrote the article as part of a special series of inaugural articles by members of the National Academy of Sciences (NAS). Elected a NAS member in 2026, he is a UC Davis Distinguished Emeritus Professor, emeritus chair of the UC Davis entomology department and emeritus provost of ASU.

In discussing the significance of the research this week, Page commented: “In this paper, we wanted to view gene expression networks from 10,000 meters and focus on the topological structure of them rather than the individual gene-gene interaction structures. I think this gives a very different global perspective on the dynamics of the networks through development, one that isn’t often given.”

“Using fat body gene expression data across five developmental time points that span an interval of behavioral transition, we construct signed gene–gene correlation networks and quantify their macro- and mesostate properties,” they related in the research article. “We then descend to the level of the gene and construct microstate gene lists of interacting genes. Behavioral maturation of worker honey bees is accompanied by coordinated, nonmonotonic changes in global network structure rather than simple global shifts in interaction strength across the network.”

“The intricate symphony of gene expression that accompanies the development of bees and their transition from nurses to foragers reveals some interesting patterns that we analyzed using tools from network science and statistical learning,” Bonabeau commented in a post on LinkedIn. “It involves a fundamental reorganization of gene networks, rather than simply changes in the expression of individual genes.”

“The implication is that behavioral differentiation may reflect collective changes in molecular organization, not just individual gene activity,” Bonabeau wrote. “Big caveat: the study demonstrates correlations, not causal regulatory mechanisms or dynamical attractors.”

Among their findings early gene networks exhibit predominantly positive correlations. Between days 6 and 10, their organization changes dramatically. Two groups of genes become internally coordinated but negatively correlated with each other: a nurse-associated, high-vitellogenin (VG) module and a forager-associated, low-VG module. --VG is an important multifunctional protein. By day 15, bees exhibit two distinct gene-expression profiles, suggesting alternative physiological states. 

Page, known for his research on honey bee behavior and population genetics, particularly the evolution of complex social behavior, constructed the first genomic map of the honey bee and pioneered the use of modern techniques to study the genetic basis of social behavior evolution in honey bees and other social insects. 

A UC Davis doctoral alumnus (1980), Page worked closely with Professor Harry H. Laidlaw Jr. , for whom the UC Davis bee biology facility is named. Together they published many significant research papers and the landmark book, “Queen Rearing and Bee Breeding” (Wicwas Press, 1998), considered the most important resource book for honey bee genetics, breeding and queen rearing.

Page maintained a UC Davis honey bee-breeding program, managed by bee breeder-geneticist Kim Fondrk, for 24 years, from 1989 to 2015. 

Page is the author of several books, “Honey Bee Genetics and Breeding (Wicwas Press, 2025); “The Art of the Bee” (Oxford University Press, 2020) and “The Spirit of the Hive” (Harvard University Press 2013). In 2023, he launched a YouTube Channel, https://youtube.com/@artofthebee.