If you ask honey bee geneticist Robert E. Page Jr. what his favorite honey is, he'll point to his wife, Michelle.

She--and any others near them--will smile every time!

Fact is, Rob Page is our favorite honey bee geneticist, and he was just named the recipient of the 2023 C. W. Woodworth Award, the highest honor accorded by the Pacific Branch, Entomological Society of America (PBESA). 

It's an honor well deserved. And a honey of an award.

“Dr. Page 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. 

Page is the 12th UC Davis recipient of the award, first presented in 1969. His mentor, and later colleague, Harry H. Laidlaw Jr. (for whom the UC Davis bee biology research facility is named), won the award in 1981.

The PBESA awards ceremony will take place at its meeting, April 2-5, in Seattle. The organization encompasses 11 Western states, and parts of Canada, Mexico and U.S. territories. 

“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 related. “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, Rob holds a bachelor's degree in entomology (1976), with a minor in chemistry, from San Jose State University. He obtained his doctorate in entomology (1980) from UC Davis; joined the UC Davis faculty in 1989; and chaired the Department of Entomology from 1999 to 2004. He gained UC Davis emeritus professor-chair status in 2004,  the year that Arizona State University (ASU) recruited him to be the founding director of its School of Life Sciences. Page organized three departments—biology, microbiology and botany, totaling more than 600 faculty, graduate students, postdoctoral fellows and staff--into one unified school. Using his UC Davis experiences, he established the school as a platform for discovery in the biomedical, genomic and evolutionary and environmental sciences; and established ASU's world-class Social Insect Research Group and Honey Bee Research Facility.

Page's career at ASU led to a series of top-level administrative roles: founding director, School of Life Sciences (2004-2010), vice provost and dean, College of Liberal Arts and Sciences (2011-2013) and university provost, 2014-2015.

Nadler praised Page's strategic vision, his leadership and his contributions to science.  He built two modern apicultural labs (in Ohio and Arizona), major legacies that are centers of honey bee research and training. The Social Insect Research Group (SIRG) at ASU is regarded as “the best in the world,” according to the late E. O. Wilson. ASU Professor Bert Hoelldobler, in an ASU news release, declared Dr. Page  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.”

Page is an Institute for Scientific Information (ISI) highly-cited author in plant and animal science. In all, he has published more than 250 research papers and articles, edited or authored five books, and is listed as a “highly-cited author” by the ISI Web of Knowledge, representing the top 1/2 of one percent of publishing scientists. Statistics include: H-index = 80, and 21,925 citations.  Since retiring from UC Davis, Dr. Page has published 65 research papers, 8 major reviews and 2 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). 

While at UC Davis, Page worked closely with Harry H. Laidlaw Jr., the father of honey bee genetics, and together they published many significant research papers and the landmark book, “Queen Rearing and Bee Breeding” (Wicwas Press, 1998). It is considered the most important resource book for honey bee genetics, breeding, and queen rearing.  Page is now in the process of updating it.

For 24 years, from 1989 to 2015, Page maintained a 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.

A 2012 Fellow of the Entomological Society of America, Page has held national and international offices. He served as secretary, chair-elect, chair, subsection cb (apiculture and social insects) of ESA from 1986-1989; president of the North American Section, International Union for the Study of Social Insects, 1991; and a Council member, International Bee Research Association, 1995-2000.

Among his 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
• Thomas and Nina Leigh Distinguished Alumni Award from UC Davis Department of Entomology and Nematology
• James Creasman Award of Excellence (ASU Alumni Association)
• UC Davis Distinguished Emeritus Professor, one awarded annually
• Distinguished Emeritus Award, College of Agricultural and Environmental Sciences, one awarded annually

In his letter of support, colleague and research collaborator James R. Carey, distinguished professor of entomology at UC Davis,  described Page as "one of the most gifted scientists, administrators, and teachers I have had the privilege to know in my 42 years in academia.”

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. Cobey, who has based her career on the Page-Laidlaw Closed Population Breeding, wrote that: “The beauty of this system is that it is practical and addresses the unique challenges of honey bee stock improvement. 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.”

Nadler also noted that “Dr. Page was involved in genome mappings of bumble bees, parasitic wasps and two species of ants. His most recent work focuses on the genetic bases of individuality in honey bees; demonstrating genetic links between pollen and nectar collection, tactile and olfactory learning characteristics, and neuroendocrine function. This work provides the most detailed understanding to date of the molecular and genetic bases to task variation in a social insect colony.” 

Nadler added: "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."

Charles William Woodworth (1865-1940), is considered the founder of both the UC Berkeley and UC Davis departments of entomology.  William Harry Lange Jr., (1912-2004) was the first UC Davis recipient of the Woodworth award (1978). Other recipients: Harry Laidlaw Jr., (1907-2003), 1981; Robert Washino, 1987; Thomas Leigh (1923-1993), 1991; Harry Kaya, 1998; Charles Summers, (1941-2021), 2009; Walter Leal, 2010; Frank Zalom, 2011; James R. Carey, 2014; Thomas Scott, 2015; and Lynn Kimsey, 2020.

Joining Rob Page in the 2023 PBESA winners' circle from UC Davis: community ecologist Louie Yang, professor, UC Davis Department of Entomology and Nematology, recipient of the Distinction in Student Mentoring Award; and UC Davis student Gary Ge, of the UC Davis Research Scholars Program in Insect Biology, who won the second annual Dr. Stephen Garczynski Undergraduate Research Scholarship. 

The complete list of this year's PBESA recipients is posted here. 

Congratulations to UC Davis-trained bee scientist Robert E. Page Jr., recipient of the 2022 Exceptional Emeriti Faculty Award from the UC Davis College of Agricultural and Environmental Sciences (CA&ES). 

Known by his colleagues as “the leading honey bee geneticist in the world,” 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 major professor 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.

Page joined the UC Davis entomology faculty in 1989 and chaired the department from 1999 to 2004 when he transitioned to emeritus and was recruited by Arizona State University (ASU)  to be the founding director of its School of Life Sciences. His career at ASU led to a series of top-level administrative roles: from founding director of the School of Life Sciences (2004-2010) to vice provost and dean, College of Liberal Arts and Sciences (2011-2013) and then to University Provost, 2014-2015. 

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).

See more about Rob Page's career on the UC Davis Department of Entomology and Nematology website.

Others honored at the CA&ES Award of Distinction ceremony will be Alumnus of the Year, Marko Zaninovich (Class of 1964); Distinguished Friends of the College, William 'Bill' Patterson and his wife, Doris Brown,  for their contributions and support of the Bohart Museum of Entomology; Exceptional Faculty Award: Daniel Sumner, the Frank H. Buck, Jr. Distinguished Professor of Agricultural and Resource Economics; and Exceptional Staff Award, Shannon Tanguay, budget fiscal officer with CA&ES. (See CA&ES website)

If you're a honey bee geneticist and you're asked to highlight your three-decade scientific career, how do you start?

Societies to genes? And how do you get from there to here?

Noted honey bee geneticist Robert E. Page Jr., a UC Davis and Arizona State University emeritus professor and administrator, has authored a newly published, invited article in the journal Geneticson “Societies to Genes: Can We Get There from Here?” that highlights his three-decade scientific career.   

 “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.

In his article, he mentions: "I studied the behavioral genetics of pollen storage in honey bees for more than 30 years. The effort was collective with my technician and colleague Kim Fondrk and numerous students and postdoctoral researchers. We used a forward genetic approach and employed bidirectional, human-assisted selection that resulted in the establishment of two strains that varied in their expression of a social phenotype. We focused on just one trait, the amount of pollen stored in the nest, a social trait that is a consequence of the interactions of thousands of bees. There are about 10–40 thousand worker bees in a honey bee colony, depending on the time of year. Honey bee colonies have a reproductive division of labor where the single queen normally lays the eggs while the workers are facultatively sterile."

"They also have a division of labor among the workers," Page wrote. "Roughly half of them are foragers, a behavior that begins when an individual is about 2–3 weeks old. Individual foragers primarily collect pollen (protein) and nectar (carbohydrate). They specialize by biasing their foraging behavior for one or the other. Specialization is partly determined by their genotype. Roughly 1/3rd of the foragers collect pollen, depending on resource availability and time of year. Pollen is collected from flowers and stored in cells of wax comb near the 'nursery,' the central part of the nest where eggs are laid in cells and larvae develop into adults. A full-sized colony may contain 10,000 larvae. Pollen is stored in excess of immediate need, it is hoarded. Some bees in the nest consume the pollen in the cells, convert the pollen proteins into vitellogenin (VG) and other proteins in the fat body that then enter the hemolymph, travel to the hypopharyngeal glands, and are converted to glandular secretions that are fed to larvae."

"Young larvae produce pheromones (chemical signals) that stimulate some of the foragers to collect pollen (Traynor et al. 2015)," he explained. "The amount of pollen stored, the number of cells that are full, affects the number of larvae that are raised and inhibits pollen foraging, thus stored pollen is regulated (Fewell and Winston 1992). Returning pollen foragers perform recruitment dances that communicate the distance and direction from the nest to their pollen sources. Other bees attend the dances and are recruited. So, the amount of stored pollen is dependent on the interactions of thousands of individual adults and larvae and is not a phenotypic trait of any individual—it is a social phenotype." 

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.

If a queen bee were to celebrate Mother's Day (and she won't because she's too busy laying eggs), what a crowded festivity that would be. 

Bee breeder-geneticist Susan Cobey of Washington State University, former manager of the Harry H.Laidlaw Jr. Honey Bee Research Facility at UC Davis, describes the queen as an "egg-laying machine."

"She's the mother of all the bees in the hive," saysCobey, who studied at UC Davis with Harry Hyde Laidlaw Jr., (1907-2003),  "the father of honey bee genetics."  During the peak season, the queen can lay up to 2000 eggs a day. That amounts to about 50,000 to 80,000 workers (sterile females) and 1000 to 2000 drones (males) in the hive. 

On her maiden flight, the queen bee mates with some 12 to 25 drones in mid-air and then she heads back to the hive to lay eggs for the rest of her life, says Cobey, internationally renowned for her Carniolan bees and classes on instrumental insemination and bee breeding (stock improvement). 

Yes, every bee in the hive has the same mother.  Not so with the fathers.

In his book, The Honey Bee Hobbyist, the Care and Keeping of Bees, Norman "Norm" Gary, UC Davis emeritus professor of apiculture, writes: "All bees in a colony develop from eggs laid by the queen, so they all share the same mother. All bees in a colony develop from eggs laid by the queen, so they all share the same mother. But as a population, they typically have around 15 fathers."

As Gary points out: "The queen bee has no control over the drones that inseminate her. (The virgin queen) mates while flying, never inside the hive." 

It's a matriarchal society. The girls (worker bees) do all the work; they serve as nurse maids, nannies, royal attendants, builders, architects, foragers, dancers, honey tenders, pollen packers, propolis or "glue" specialists, air conditioning and heating technicians, guards, and undertakers. So their abbreviated life (during the summer the life span of a worker bee is only four to six weeks) is not surprising. The drones, or males, serve only a reproductive function. Once they they mate, they die.

Honey bee geneticist Robert E. Page Jr., distinguished emeritus professor of entomology at UC Davis (and emeritus provost, Arizona State University) studied with Laidlaw for his doctorate at UC Davis. He pays tribute to Laidlaw  in his book,  The Art of the Bee: Shaping the Environment from Landscapes to Societies.

In Chapter Nine, "The Song of the Queen," Page reprinted a poem by E. B. White (Dec. 15, The New Yorker) objecting to instrumental insemination. White opined in the poem that the queen bee should "mate with whatever drone" she encounters. 

Page reprinted Laidlaw's response, published in the San Francisco Chronicle, which said in part:

Her offspring slave throughout the day,
They feed her children as best they may
They would like to see a moment
Directed toward stock improvement. 

If you're interested in bees and beekeeping, or just curious about these amazing superorganisms, these books read well on Mother's Day...and any other day.