- Author: Kathy Keatley Garvey
He will be honored at the CA&ES Award of Distinction dinner and celebration awards ceremony on Thursday, Nov. 3 in the UC Davis Activities and Recreation Center Ballroom.
Page, emeritus professor and chair of the UC Davis Department of Entomology and an emeritus professor and administrator at Arizona State University, “is a pioneering researcher in the field of evolutionary genetics and social behavior of honey bees, and a highly respected and quoted author, teacher and former administrator,” wrote nominator Steve Nadler, professor and chair of the UC Davis Department of Entomology and Nematology.
“One of Dr. Page's most salient contributions to science was to construct the first genomic map of the honey bee, which sparked a variety of pioneering contributions not only to insect biology but to genetics at large,” Nadler pointed out. “It was the first genetic map of any social insect. He was the first to demonstrate that a significant amount of observed behavioral variation among honey bee workers is due to genotypic variation. In the 1990s, he and his students and colleagues isolated, characterized and validated the complementary sex determination gene of the honey bee; considered the most important paper yet published about the genetics of Hymenoptera. The journal Cell featured their work on its cover. In subsequent studies, he and his team published further research into the regulation of honey bee foraging, defensive and alarm behavior.”
A native of Bakersfield, Kern County, Rob holds a bachelor's degree in entomology (1976), with a minor in chemistry, from San Jose State University. He received his doctorate in entomology in 1980 from UC Davis, where he studied with his doctoral research mentor, Harry H. Laidlaw Jr., “the father of honey bee genetics” for whom the university's bee facility, the Harry H. Laidlaw Jr. Honey Bee Research Facility on Bee Biology Road, is named.
While at UC Davis, Page worked closely with Laidlaw. 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.
For 24 years, from 1989 to 2015, Page maintained a UC Davis honey bee-breeding program, managed by bee breeder-geneticist Kim Fondrk. Their contributions include discovering a link between social behavior and maternal traits in bees. Their work was featured in a cover story in the journal Nature. In all, Nature featured his work on four covers from work mostly done at UC Davis.
Since his retirement from UC Davis, Page has published 65 research papers, eight major reviews and two scholarly books, many using his UC Davis affiliation. He authored “The Spirit of the Hive: The Mechanisms of Social Evolution” (Harvard University Press, 2013) and the “Art of the Bee: Shaping the Environment from Landscapes to Societies” (Oxford University Press, 2020).
Now residing near Davis, Page continues to focus his research on honey bee behavior and population genetics, particularly the evolution of complex social behavior. His continuing academic activities bring credit to bee biology and UC Davis, Nadler said. “His large number of publications and citations continue to be an important component of the high national rating of our entomology department.”
Page continues to work closely with UC Davis professors and students, offering advice, helping them with grants, and editing manuscripts. A few years ago, he held an international workshop at the Laidlaw facility. He teaches courses (including “The Social Contract: from Rousseau to the Honey Bee,” and “The Song of the Queen: Thrilling Tales of Honey Bee Mating Behavior”) for the UC Davis Osher Lifelong Learning Institute (OLLI).
“Not surprisingly, Dr. Page humbly considers his most far-reaching and important accomplishment, the success of his mentees, including at least 25 graduate students and postdocs who are now faculty members at leading research institutions around the world,” Nadler wrote. “He also built two modern apicultural labs (in Ohio and Arizona), major legacies that are centers of honey bee research and training. He has trained many hundreds of beekeepers. His public service now extends to working as a Fellow with the California Academy of Sciences.”
Page's colleagues praise his strategic vision, his leadership and his contributions to science. ASU Professor Bert Hoelldobler described him as “the leading honey bee geneticist in the world. A number of now well-known scientists in the U.S. and Europe learned the ropes of sociogenetics in Rob's laboratory.”
Bee breeder-geneticist Susan Cobey of Washington State University, former manager of the Laidlaw facility, emphasized Page's importance to the bee breeding and beekeeping industry. She has based her entire career on the Page-Laidlaw Closed Population Breeding. “The beauty of this system is that it is practical and addresses the unique challenges of honey bee stock improvement,” she related. “Queens mate in flight with numerous drones and selection is based upon complex behaviors at the colony level, influenced by the environmental. Hence, traditional animal breeding models do not apply well to honey bees.”
When Page addressed the 2016 meeting of the International Congress of Entomology (ICE), he was introduced as “an engaging, energetic, humorous and highly intellectual speaker; this will be a memorable, entertaining and highly educational lecture by one of the most important entomologists of the 21stt century. His background is in behavior and population genetics, and the focus of his current research is on the evolution of complex social behavior. Using the honey bee as a model, Dr. Page has dissected their complex foraging division of labor at all levels of biological organization, from gene networks to complex social interactions.”
Among Page's many honors:
- Fellow of the American Association for the Advancement of Science
- Awardee of the Alexander von Humboldt Senior Scientist Award (the Humboldt Prize - the highest honor given by the German government to foreign scientists)
- Foreign Member of the Brazilian Academy of Sciences
- Fellow of the American Academy of Arts and Sciences
- Elected to the Leopoldina, the German National Academy of Sciences (the longest continuing academy in the world)
- Fellow of the Wissenschaftskolleg zu Berlin
- Fellow of the Entomological Society of America
- Awardee of the Carl Friedrich von Siemens Fellowship
- Fellow of the California Academy of Sciences
- Fellow, Carl Friedrich von Siemens Foundation, Munich, Germany, September 2017-August 2018
- Thomas and Nina Leigh Distinguished Alumni Award from UC Davis Department of Entomology and Nematology
- James Creasman Award of Excellence (ASU Alumni Association)
- Regents Professor, Arizona State University
- UC Davis Distinguished Emeritus Professor
![Noted honey bee geneticist Robert E. Page Jr. checks out a bee swarm at Arizona State University. Noted honey bee geneticist Robert E. Page Jr. checks out a bee swarm at Arizona State University.](https://ucanr.edu/blogs/entomology/blogfiles/95610.jpg)
![Former UC Davis graduate student Robert E. Page Jr., with doctoral research mentor, Harry Hyde Laidlaw Jr. 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. Former UC Davis graduate student Robert E. Page Jr., with doctoral research mentor, Harry Hyde Laidlaw Jr. 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.](https://ucanr.edu/blogs/entomology/blogfiles/95612.jpg)
- Author: Kathy Keatley Garvey
Honey bee geneticist and UC Davis alumnus Robert E. Page Jr., will teach a course on "The Social Contract: from Rousseau to the Honey Bee" on Monday, Feb. 7 from 2 to 4 p.m. He is an emeritus professor and former chair of the UC Davis Department of Entomology and Nematology and a university provost emeritus and a regents' professor emeritus at Arizona State University. A UC Davis alumnus and the 2019 recipient of the UC Davis Distinguished Emeritus Award, administered by the UC Davis Emeriti Association, Page has published more than 230 research papers and three books on honey bee genetics and behavior. Course specifics: 213SNR388 $25 Zoom.
Molecular geneticist and physiologist Joanna Chiu, professor and vice chair of the UC Davis Department of Entomology and Nematology and a Chancellor's Fellow, will cover "The Circadian Clock and Chronomedicine" on Wednesday, March 23 from 10 a.m. to noon. Chiu, who holds a doctorate in molecular genetics from New York University, received postdoctoral training at the Center for Advanced Biotechnology and Medicine at Rutgers University. Course specifics: 213SNR383 $25 Zoom.
Social Contract: from Rousseau to the Honey Bee Animals
"Animals that live together in a society, like the social insects, have a tacit agreement, a social contract, that guarantees that their interests are protected in exchange for their social cooperation," Page says. "Like social contracts that bind human societies, this contract isn't written on paper, it is implied, though in humans its enforcement is expressed in explicit written laws and national constitutions. The social contract of insects has been written by natural selection in their DNA over thousands of generations. In this class we will explore elements of the social contract of honey bee societies, its origins and 'laws for enforcement,' and the social services honey bees get as a consequence of being members of a society. Specifically, we will explore their systems of national defense, internal police, public works, public health and border control. Without these features, their society would fail, as would ours. Discover what we can learn about ourselves from studying social insects."
"The circadian clock is an internal body clock that controls all aspects of physiology and behavior in every single one of us. It interprets environmental and metabolic time cues to ensure our body performs at its best," Chiu says. "A sequel to the OLLI course 'What Time is it and Why Does it Matter?,' this course will revisit the very important, yet infrequently discussed physiological machinery that controls sleep- wake cycles, hormone production and immune system. Because disruptions in the circadian clock have been associated with a range of human diseases, we will highlight life-style choices and work schedules that cause clock disruptions. Finally, we will discuss new strategies in the medical field that leverage circadian biology concepts to increase the efficacy of medical treatments."
OLLI is headquartered at 463 California Ave. Davis, CA 95616. Members of the UC Davis Retirees' Association and the UC Davis Emeriti Association are offered a complimentary course credit equivalent to their paid OLLI membership fee.
The annual OLLI membership is $60 for the academic year (Oct. 4, 2021- June 30, 2022). More information on OLLI, which is part of the UC Davis Continuing and Professional Education, is available online or contact (530) 752-9695 or olli@ucdavis.edu
![Honey bee geneticist Robert E. Page Jr. examines a swarm. Honey bee geneticist Robert E. Page Jr. examines a swarm.](https://ucanr.edu/blogs/entomology/blogfiles/87651.jpg)
- Author: Kathy Keatley Garvey
“I was thrilled to be invited to write this perspectives/review of my scientific career; it is a collection of 30 years of single-minded focus on one question,” said Page, who is renowned for his research on honey bee behavior and population genetics, particularly the evolution of complex social behavior, and for his work on the first genomic map of the honey bee.
"The editors contacted me to write a perspectives/review that focuses on my own work, a study in complex adaptation,” Page related. “This is the first time they have done a perspectives article like this. I was, of course, honored. More than half of the work was done at UC Davis.”
“Understanding the organization and evolution of social complexity is a major task because it requires building an understanding of mechanisms operating at different levels of biological organization from genes to social interactions,” Page wrote in his abstract. “I discuss here, a unique forward genetic approach spanning more than 30 years beginning with human-assisted colony-level selection for a single social trait, the amount of pollen honey bees (Apis mellifera L.) store. The goal was to understand a complex social trait from the social phenotype to genes responsible for observed trait variation.”
“The approach,” Page wrote, “combined the results of colony-level selection with detailed studies of individual behavior and physiology resulting in a mapped, integrated phenotypic architecture composed of correlative relationships between traits spanning anatomy, physiology, sensory response systems, and individual behavior that affect individual foraging decisions. Colony-level selection reverse engineered the architecture of an integrated phenotype of individuals resulting in changes in the social trait. Quantitative trait locus (QTL) studies combined with an exceptionally high recombination rate (60 kb/cM), and a phenotypic map, provided a genotype–phenotype map of high complexity demonstrating broad QTL pleiotropy, epistasis, and epistatic pleiotropy suggesting that gene pleiotropy or tight linkage of genes within QTL integrated the phenotype. Gene expression and knockdown of identified positional candidates revealed genes affecting foraging behavior and confirmed one pleiotropic gene, a tyramine receptor, as a target for colony-level selection that was under selection in two different tissues in two different life stages. The approach presented here has resulted in a comprehensive understanding of the structure and evolution of honey bee social organization.”
Page, who received his doctorate in entomology (1980) from UC Davis, joined the UC Davis entomology faculty in 1989, and chaired the department from 1999-2004. In 2004, Arizona State University recruited him as founding director of its School of Life Sciences. His career advanced from dean of Life Sciences, to vice provost and dean of the College of Liberal Arts and Sciences, to university provost. Today he holds the titles of ASU provost emeritus, ASU Regents professor emeritus, and UC Davis distinguished emeritus professor, an award bestowed in 2019.
At UC Davis, Page worked closely with Harry H. Laidlaw Jr., (the father of honey bee genetics) for whom the university's bee facility is named.
For 24 years, from 1989 to 2015, Page maintained a UC Davis honey bee-breeding program, managed by bee breeder-geneticist Kim Fondrk at the Harry H. Laidlaw Jr. Honey Bee Research Facility. Together they discovered a link between social behavior and maternal traits in bees.
Page has authored than 250 research papers, including five books: among them, The Spirit of the Hive: The Mechanisms of Social Evolution, published by Harvard University Press in 2013. His most recent book is The Art of the Bee: Shaping the Environment from Landscapes to Societies, published by Oxford University Press 2020. (See news release on why bees are both artists and engineers.)
Page is a highly cited author on such topics as Africanized bees, genetics and evolution of social organization, sex determination, and division of labor in insect societies.
![](https://ucanr.edu/blogs/entomology/blogfiles/85358.jpg)
![Honey bee geneticist Robert E. Page Jr. examines a bee swarm. He recently wrote a Genetics journal article, Honey bee geneticist Robert E. Page Jr. examines a bee swarm. He recently wrote a Genetics journal article,](https://ucanr.edu/blogs/entomology/blogfiles/85360.jpg)
- Author: Kathy Keatley Garvey
As environmental artists, bees are "responsible for the brilliantly colored flowers in our landscapes," and as environmental engineers, they engineer “the niches of multitudes of plants, animals and microbes.”
Page, with UC Davis roots and Arizona State University wings, has just authored a 256-page book, “The Art of the Bee: Shaping the Environment from Landscapes to Societies” (Oxford University Press), to be published Aug. 6.
“It's a long time in the making,” said Page, who received his doctorate in entomology at UC Davis and served as a professor and chair of the Department of Entomology (now Entomology and Nematology) before heading to Arizona State University (ASU), where he advanced to school director, college dean and university provost.
“Twenty-five years ago, my friend and mentor Harry Laidlaw (for whom the UC Davis bee facility is named) wanted to write a honey bee biology textbook,” Page recalled. When they finished the outline, “it looked very much like the excellent book by Mark Winston The Biology of the Honey Bee, published in 1987 by Harvard University Press. I decided we didn't need another one, and we still don't.”
The book differs in that it's a collection of “sparkling essays” that “read like mystery stories,” said Rudiger Wehner, professor and director emeritus of the Institute of Zoology, University of Zürich. “With these lucidly written stories, Page takes us on a delightful journey through the many biological traits that on the whole constitute the honeybees' social contract.”
“But don't be fooled by the amiable and personal style—the book is comprehensive—from colony collapse disorder to colony-level evolution—and chock full of the latest results, presented with clarity and depth, leavened with razor-sharp insights into social evolution,” noted Gene Robinson, director, Carl R. Woese Institute for Genomic Biology and Department of Entomology, University of Illinois at Urbana-Champaign.
Page said his book is geared toward “the person who has a basic knowledge of biology and a fascination with bees, perhaps an educated hobby beekeeper--there are a lot of them--or an undergraduate or graduate student with an interest.”
In addition to chapters on environmental artists and environmental engineering, Page includes chapters on social contracts, superorganisms, reproductive competitions, and concludes with “The song of the queen.”
In the epilogue, Page ponders the complexity of individual bees and their colonies, comparing them to humans. “Members of complex societies live close together in closed nests, shared home sites, villages, etc., or in closely connected nomadic tribes. As groups, they typically have a set of tacit rules by which they live that involves working for the good of the group, systems of group and resource defense, internal mechanisms of policing cheaters that don't cooperate and live by the rules, a division of labor often associated with group defense and gathering and sharing resources, and usually asymmetries and rules associated with reproduction. These same general characteristics seem to apply broadly across eusocial insects (aphids, termites, bees, ants, and wasps), eusocial rodents (naked mole rats), higher apes, and humans. Why? The similarities are inescapable due to the nature of social contracts; they must have specific elements to protect the power and will of individuals, whether citizens of the United States of America or workers in a honey bee colony. The contract binds individuals to a society, but the specific social organization evolves by reverse engineering. Natural selection acts on the whole colony; social structure evolves to fit the needs of the group within a given environment. “
Page points out that “Anthropocentric thinking can obscure the way we view nature and lead to false conclusions. Look at Aristotle and honey bee division of labor: For more than 2,000 years it was thought that the bees that work in the nest were postpubescent old men because they're hairy! In fact, the older bees forage and aren't hairy because the hairs break off as they age. I now see my work in a new light; we aren't so different, bees and humans. The elements of our social structures, and how they come about, have many similarities.”
Page is known for his research on honey bee behavior and population genetics, particularly the evolution of complex social behavior. One of his most salient contributions to science was to construct the first genomic map of the honey bee, which sparked a variety of pioneering contributions not only to insect biology but to genetics at large.
At UC Davis, he maintained a honey bee-breeding program for 24 years, from 1989 to 2015, managed by bee breeder-geneticist Kim Fondrk at the Harry H. Laidlaw Jr. Honey Bee Research Facility. They discovered a link between social behavior and maternal traits in bees.
UC Davis named him the 2019 distinguished emeritus professor. Nominator Steve Nadler, professor and chair of the UC Davis Department of Entomology and Nematology, praised Page as “a pioneer researcher in the field of behavioral genetics, an internationally recognized scholar, a highly respected author, a talented and innovative administrator, and a skilled teacher responsible for mentoring many of today's top bee scientists…he is arguably the most influential honey bee biologist of the past 30 years.”
Page has authored more than 250 research papers, including five books. Among them “The Spirit of the Hive: The Mechanisms of Social Evolution” (Harvard University Press, 2013) and “Queen Rearing and Bee Breeding,” with Harry H. Laidlaw (Wicwas Press, 1997). He is a highly cited author on such topics as Africanized bees, genetics and evolution of social organization, sex determination, and division of labor in insect societies.
Page, who received his doctorate in entomology from UC Davis in 1980, joined the UC Davis faculty in 1989 and left as emeritus chair of the Department of Entomology in 2004 when ASU recruited him for what would become a series of top-level administrative roles. He advanced from director of the School of Life Sciences to dean of Life Sciences; vice provost and dean of the College of Liberal Arts and Sciences; and university provost. Today he holds the titles of provost emeritus of ASU and Regents professor emeritus, as well as UC Davis department chair emeritus, professor emeritus, and UC Davis distinguished emeritus professor.
Page is an elected member of the American Academy of Arts and Sciences, the Brazilian Academy of Science, Leopoldina (the German National Academic of Science), and the California Academy of Science. He is a recipient of the Alexander von Humboldt Senior Scientist Award (Humboldt Prize, 1995), the Carl Friedrich von Siemens Fellowship (2013), James W. Creasman Award of Excellence at ASU (2018).
![Honey bee geneticist and biologist Robert E. Page Jr. with his book, Honey bee geneticist and biologist Robert E. Page Jr. with his book,](https://ucanr.edu/blogs/entomology/blogfiles/72700.jpg)
![Honey bee geneticist and biologist Robert E. Page Jr., author of Honey bee geneticist and biologist Robert E. Page Jr., author of](https://ucanr.edu/blogs/entomology/blogfiles/72701.jpg)
- Author: Kathy Keatley Garvey
His talk, free and open to the public, is sponsored by the UC Davis Emeriti Association and the UC Davis Retirees' Association.
“Honey bees exhibit complex social behavior that rivals our own,” Page says. “It is fundamentally bound within a social contract much like ours that makes the basic social structure inescapable, a consequence of living together in family groups. Social structures evolve by natural selection operating on the final product, the colony as a reproductive unit. The structures themselves are reverse engineered.”
“I will show how selection on the economy of the colony shapes structures from nest and social architecture to gene networks.”
Page is a UC Davis Distinguished Emeritus Professor, an honor awarded in January; and a Regents Professor Emeritus and University Provost Emeritus from Arizona State University, Tempe. He retired from UC Davis in May 2004 after chairing the Department of Entomology, now the Department of Entomology and Nematology.
Born and reared in Bakersfield, Kern County, Rob received his bachelor's degree in entomology, with a minor in chemistry, from San Jose State University in 1976. After obtaining his doctorate from UC Davis in 1980, he served as assistant professor at The Ohio State University before joining the UC Davis entomology faculty in 1989 as an associate professor. He began working closely with Harry H. Laidlaw Jr., (the father of honey bee genetics) for whom the university's bee facility is named. Together they published many significant research papers.
Page chaired the Department of Entomology from 1999 to 2004, when Arizona State University recruited him to be the founding director of the School of Life Sciences of Arizona State University (ASU). His ASU career advanced to dean of Life Sciences; vice provost and dean of the College of Liberal Arts and Sciences; and university provost.
Page is known for his research on honey bee behavior and population genetics, particularly the evolution of complex social behavior. One of his most salient contributions to science was to construct the first genomic map of the honey bee, which sparked a variety of pioneering contributions not only to insect biology but to genetics at large.
At UC Davis, he maintained a honey bee-breeding program for 24 years, from 1989 to 2015, managed by bee breeder-geneticist Kim Fondrk at the Harry H. Laidlaw Jr. Honey Bee Research Facility. They discovered a link between social behavior and maternal traits in bees. Their work was featured in a cover story in the journal Nature. In all, Nature featured his work on four covers from work mostly done at UC Davis.
Page and his lab pioneered the use of modern techniques to study the genetic basis of social behavior evolution in honey bees and other social insects. He was the first to employ molecular markers to study polyandry and patterns of sperm use in honey bees. He provided the first quantitative demonstration of low genetic relatedness in a highly eusocial species.
His work has garnered a significant impact in the scientific community through his research on the evolutionary genetics and social behavior of honey bees. He was the first to demonstrate that a significant amount of observed behavioral variation among honey bee workers is due to genotypic variation. In the 1990s, he and his students and colleagues isolated, characterized and validated the complementary sex determination gene of the honey bee; considered the most important paper yet published about the genetics of Hymenoptera. The journal Cell featured their work on its cover. In subsequent studies, he and his team published further research into the regulation of honey bee foraging, defensive and alarm behavior.
Page has authored than 250 research papers, including five books: among them “The Spirit of the Hive: The Mechanisms of Social Evolution,” published by Harvard University Press in 2013. He is a highly cited author on such topics as Africanized bees, genetics and evolution of social organization, sex determination, and division of labor in insect societies. His resume shows more than 18,000 citations.
Highly honored by his peers, Page is a fellow of a number of organizations, including the American Association for the Advancement of Science, the California Academy of Sciences, the Entomological Society of America, and organizations in Germany and Brazil. He received the Alexander von Humboldt Senior Scientist Award, known at the Humboldt Prize, the highest honor given by the German government to foreign scientists. He received the 2018 Thomas and Nina Leigh Distinguished Alumni Award from UC Davis Department of Entomology and Nematology.
The UC Davis Retiree Center periodically holds noon educational seminars ("BrainFood Talks") featuring timely speakers. For more information, contact the UC Davis Retiree Center at (530) 752-5182 or retireecenter@ucdavis.edu.
![Noted honey bee geneticist checking out a bee swarm at the University of Arizona, Tempe. Noted honey bee geneticist checking out a bee swarm at the University of Arizona, Tempe.](https://ucanr.edu/blogs/entomology/blogfiles/64199.jpg)