- Author: Kathy Keatley Garvey
Page, a UC Davis doctoral alumnus, a UC Davis distinguished emeritus professor and former chair of the UC Davis Department of Entomology, wrote an article The Art of the Bee for the journal, and editor Robert Brodschneider interviewed him in a piece titled Robert E. Page Jr--Mapper of the Genetic Architecture of the Honey Bee.
Rob lives and breathes bees, UC Davis, and Arizona State University (ASU). He obtained his doctorate in entomology in 1980 from UC Davis; joined the UC Davis faculty in 1989; and chaired the Department of Entomology from 1999 to 2004. After retiring from UC Davis in 2004, he accepted an appointment at ASU as founding director of the School of Life Sciences. He served as provost of ASU from 2013- 2015, and dean of the College of Liberal Arts and Sciences, 2011-2013.
His research on honey bee behavior and genetics appears in his publications: Queen Rearing and Bee Breeding (1997, with Harry H. Laidlaw Jr., his major professor at UC Davis and "the father of honey bee genetics"); The Spirit of the Hive, Harvard University Press (2013); and The Art of the Bee, Oxford University Press (2020). His 230-plus research papers have been cited more than 20,000 times.
Much of Rob Page's research occurred at UC Davis. For 24 years, from 1989 to 2015, he maintained a honey bee-breeding program, managed by bee breeder-geneticist Kim Fondrk, at the Harry H. Laidlaw Jr. Honey Bee Research Facility on Bee Biology Road, west of the central campus. 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.
Looking to learn more about bees? You'll want to pick up a copy of his book, ;The Art of the Bee: Shaping the Environment from Landscapes to Societies, 2020, Oxford University Press), and/or access his newly created free YouTube channel, The Art of the Bee," at https://www.youtube.com/@artofthebee.
Why did Page create the free and accessible-to-all YouTube Channel? Because that's what Alexander von Humboldt (1769-1859), known as a German geographer, naturalist, explorer, and proponent of Romantic philosophy and science, would have done.
It's about making science understandable.
"Von Humboldt appealed to artists to learn about nature, and ecology, and paint it," Page wrote in the Bee World article. "He believed that artists and writers could do more to advance an understanding of science and nature than the scientific specialist. His plea was for making science understandable to the public, a plea for popular science."
Page's YouTube channel guides the viewer through "the fascinating biology and behavior of the bee," presented in 38 videos ranging from 4 to 27 minutes in length. He organized the videos into six segments that roughly correspond to the nine chapters of the book:
- Landscape Artists
- Environmental Engineers
- The Social Contract
- Superorganisms
- How to Make a Superorganism
- Song of the Queen.
We love this passage from his book:
"The impact of bees on our world is immeasurable. Bees are responsible for the evolution of the vast array of brightly-colored flowers and for engineering the niches of multitudes of plants, animals, and microbes. They've painted our landscapes with flowers through their pollination activities and have evolved the most complex societies to aid their exploitation of the environment."
We can learn so much from the bees...
- Author: Kathy Keatley Garvey
The link: https://youtube.com/@artofthebee
Just call it "the fascinating world of bees!"
The content is from his book, The Art of the Bee: Shaping the Environment from Landscapes to Societies, (Oxford University Press, 2020).
"The impact of bees on our planet is immeasurable," he says. "Bees are responsible for the evolution of the vast array of brightly-coloured flowers and for engineering the niches of multitudes of plants, animals, and microbes. They've painted our landscapes with flowers through their pollination activities and have evolved the most complex societies to build and exploit the environment. The biology of the honey bee is one that reflects their role in transforming environments with their anatomical adaptations and a complex language that together function to harvest floral resources. A social system that includes a division of labor builds, defends, and provisions nests containing tens of thousands of individuals, only one of whom reproduces."
"This YouTube channel series presents fundamental biology, not in organizational layers, but wrapped around interesting themes and concepts, and in ways designed to explore and understand each concept," explained Page, who retired in 2019. "It examines the co-evolution of bees and flowering plants, bees as engineers of the environment, the evolution of sociality, the honey bee as a superorganism and how it evolves, and the mating behavior of the queen."
The content is divided into six segments:
Episode 1: “Darwin's Abominable Mystery”
Episode 2: “Floral Adaptations”
Episode 3: “Adaptations of Bees”
Episode 4: “Dance Language”
Episode 5: “Navigation”
Episode 6: “Time Scales of Change”
Environmental Engineers:
Episode 1: “Environmental Engineering”
Episode 2: “Niche Construction”
Episode 3: “Nest Defense in Niche Construction”
The Social Contract:
Episode 1: “Political Philosophy of Bees Social Contract”
Episode 2: “Complex Social Structures”
Episode 3: “Power and Will of Social Insects”
Episode 4: “Evolution of Altruism in Bees”
Episode 5: “Public Health and Bees”
Episode 6: “Honey Bee Public Works, Welfare Immigration”
Superorganisms:
Episode 1: “What Is a Superorganism?”
Episode 2: “Reproduction, Protection and Nutrition”
Episode 3: “Biogenic Law and Baers Law”
Episode 4: “Germ Plasm Theory”
Episode 5: “A Metaphor or an Entity”
How to Make a Superorganism:
Episode 1: “The Spirit of the Hive”
Episode 2: “Division of Labor”
Episode 3: “Colony Level Selection”
Episode 4: “Phenotypic Architecture”
Episode 5: “Phenotypic and Genetic Architectures”
Episode 6: “Bee Development”
Song of the Queen:
Episode 1: “Natural History and Castes”
Episode 2: “The Song Begins: Making a New Queen”
Episode 3: “Conditions that Stimulate Queen Rearing”
Episode 4: “Where Do Queens Mate?”
Episode 5: “How Do Queens Mate? How Many Times?”
Episode 6: “Polyandry and Sperm Use”
Page joined Arizona State University in 2004, after retiring as Professor Emeritus and Chair Emeritus, UC Davis Department of Entomology, to be founding director of the School of Life Sciences. He served as provost of Arizona State University (2013- 2015) and dean of the College of Liberal Arts and Sciences (2011-2013). His research on honey bee behavior and genetics appears in his publications Queen Rearing and Bee Breeding (1997, with Harry H. Laidlaw Jr. , his major professor at UC Davis and "the father of honey bee genetics"); The Spirit of the Hive, Harvard University Press (2013); and The Art of the Bee, Oxford University Press (2020). His 230-plus research papers have been cited more than 20,000 times.
Much of his research occurred at UC Davis. 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.
Page's UC Davis honors include:
- Thomas and Nina Leigh Distinguished Alumni Award (2018) from UC Davis Department of Entomology and Nematology
- UC Davis Distinguished Emeritus Professor (2019), one awarded annually, UC Davis Emeriti Association
- Exceptional Emeriti Faculty Award (2022), UC Davis College of Agricultural and Environmental Sciences
He most recently won the 2023 C. W. Woodworth Award, the highest honor that the Pacific Branch of ESA offers. “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. Laidlaw won the award in 1981. (See news story)
- Author: Kathy Keatley Garvey
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.
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."
"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.
- Author: Kathy Keatley Garvey
Renowned bee geneticist and biologist Robert E. Page Jr., who maintains strong ties to UC Davis and Arizona State University, clarifies this in his newly published book, The Art of the Honey Bee; Shaping the Environment from Landscapes to Societies.
"Foraging honey bees are often grouped into pollen or nectar specialists," Page writes. "However, many or most foragers collect both pollen and nectar. Some bias their foraging toward collecting nectar, others toward pollen. A bee with a strong nectar forging bias accidentally brushes against the anthers of a flower and get pollen on her body hairs while she probes the nectary. After collecting nectar, she grooms herself and stores a small amount of pollen on her hind legs. When she returns to the nest, she brings a load of nectar and a small amount of pollen."
Page points out that foragers have "loading limits." Loading limits: isn't that a wonderful description, especially when you see a honey bee weighted down with a huge ball of pollen?
"Pollen and nectar collecting are intimately related because of loading limits experienced by foragers," Page relates in his 256-page book, 25 years in the making. "A fully loaded bee collecting only nectar can carry about 60 milligrams (mg). A fully loaded pollen forager carries about 30 milligrams."
In his book, published by Oxford University Press, Page covers honey bees as environmental artists, responsible for the "brilliantly colored flowers in our landscapes," and as engineers, engineering "the niches of multitudes of plants, animals and microbes."
The book, chock-full of information, is both well-written and comprehensive. It's a "must have" for anyone who wants to learn more about bees and understand them. Page writes in an easy-going, easy-to-understand style.
“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.
The book, Page says, 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.”“
The internationally renowned bee geneticist 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 received his doctorate in entomology from UC Davis in 1980, and joined the UC Davis faculty in 1989. in 2004, when he was chair of the Department of Entomology, Arizona State University (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.
Highly honored by his peers, 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).
- Author: Kathy Keatley Garvey
And it's sure to "bee" among the very best.
Eminent honey bee geneticist and biologist Robert E. Page Jr. has 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.
Did you know that honey bees are both artists and engineers?
As environmental artists, they 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.”
The book "is 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.”
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.”
“Prior to the rise of flowering plants, the landscape was dull,” Page begins. “The first plants invaded land more than 450 million years ago. Species of ferns, horsetails, club mosses, and other “primitive,” non-seed-producing, non-flowering plants dominated. The earth was painted from a palette of green and brown. The first seed plants were gymnosperms, which originated around 100 million years later and eventually gave rise to the conifers that dominated the earth with massive forests. The first flowering plants, the angiosperms, appeared perhaps as early as 250 million years ago. The rise of flowering plants resulted in a burst of new plant species as they adapted to their insect pollinators, mostly beetles and short-tongued flies. Think of how the landscape changed. “
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.
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).