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The Spirit of the Hive
That's the title of a newly published book written by Robert E. Page Jr., one of the world's foremost honey bee geneticists.
In his 224-page book, published by Harvard University Press, Page sheds light on how 40,000 bees, "working in the dark, seemingly by instinct alone, could organize themselves to contstruct something as perfect a a honey comb."
Page, former professor and chair of the UC Davis Department of Entomology, marvels at how bees can accomplish these incredible tasks. In synthesizing the findings of decades of experiments, he presents "a comprehensive picture of the genetic and physiological mechanisms underlying the division of labor in honey bee colonies and explains how bees' complex social behavior has evolved over millions of years," according to the Harvard University flier.
Page, now vice provost and dean of the Arizona State University's College of Liberal Arts and Sciences and Foundation Professor of Life Sciences, still keeps his specialized stock of honey bees at the Harry H. Laidlaw Jr. Honey Bee Research Facility at UC Davis. Bee breeder-geneticist Michael “Kim” Fondrk, who worked with Page at Ohio State University, UC Davis and ASU, manages the stock.
In his book, Page talks about the coordinated activity of the bees and how worker bees respond to stimuli in their environment. The actions they take in turn alter the environment, Page says, and "so change the stimuli for their nestmates. For example, a bee detecting ample stores of pollen in the hive is inhibited from foraging for more, whereas detecting the presence of hungry young larvae will stimulate pollen gathering."
Division of labor, Page says, is an inevitable product of group living because "individual bees vary genetically and physiologically in their sensitivities to stimuli and have different probabilities of encountering and responding to them."
Page, who received his doctorate in entomology at UC Davis in 1980, served as an assistant professor at Ohio State University before joining the UC Davis Department of Entomology in 1989. He chaired the department for five years, from 1999 to 2004.
In 2004--the year Page retired from UC Davis--ASU recruited him as the founding director and dean of the School of Life Sciences. At the time, his duties included organizing three departments—biology, microbiology and botany, totaling more than 600 faculty, graduate students, postdoctoral fellows and staff--into one unified school.
As its founding director, Page established the school as a platform for discovery in the biomedical, genomic and evolutionary and environmental sciences. He also established ASU’s Honey Bee Research Facility.
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.
Add this one to the list: The Spirit of the Hive.
The queen and her court. (Photo by Kathy Keatley Garvey)
Queen cells. (Photo by Kathy Keatley Garvey)
Turning a Page in Entomological History
The ESA Governing Board today announced the new fellows, who will be recognized Nov. 11, 2012 at the ESA’s annual meeting in Knoxville, Tenn.
Page, the vice provost and dean of the College of Liberal Arts and Sciences, Arizona State University, Tempe, since July 2011, is a pioneer in the field of evolutionary genetics and the social behavior of honey bees.
Page did much of his research at UC Davis. He received his doctorate in entomology at UC Davis in 1980, learning from his major professor Harry H. Laidlaw Jr. (for whom the UC Davis bee research facility is named). He then joined The Ohio State University faculty before returning to UC Davis as a faculty member in 1989.
He chaired the UC Davis Department of Entomology from 1999 to 2004.
Page, who studies the evolution of complex social behavior in honey bees, from genes to societies, left UC Davis in 2004 to be the founding director of the new School of Life Sciences. Arizona State University, where he built a Social Insect Research Group that is now recognized worldwide.
Page was trained as an entomologist, evolutionary population geneticist, classical animal breeder, and mechanistic behaviorist, according to a news release issued by the ESA. "This training has defined his research approach of looking at the genetics and evolution of complex social behavior. He has taken a vertical approach to understanding the mechanisms of honey bee social foraging and how it evolves."
His work is contained in more than 225 research articles. Page has also co-edited three books and authored or co-authored two. Page is a highly cited ISI (Institute for Scientific Information) author in plant and animal science. He is a fellow of the American Association for the Advancement of Science (AAAS) and an elected member of the American Academy of Arts and Sciences, the German National Academy of Science, and the Brazilian Academy of Science. In 1995 he was awarded the Alexander von Humboldt Research Prize by the government of Germany.
Although Page resides in Arizona, his research bees are UC Davis-based residents. Bee breeder-geneticist Michael “Kim” Fondrk, who worked with Page at Ohio State University, UC Davis and ASU, manages the specialized genetic stock, which today includes 90 hives.
Of the 90 hives, about 70 have instrumentally inseminated queens as part of their pollen-hoarding breeding research program.
Back in 2006, Page told us: "Davis is still home to me. I raised my family there, my closest friends and colleagues are still in the entomology department there. I still feel very strong attachments.”
Former department chair Robert Washino remembers him fondly. He recently recalled that "Rob chose to return to UC Davis in 1989 to be with his former major professor (Harry H. Laidlaw Jr.). He later cared for him and his wife, Ruth, in their declining years. We all remember Rob’s scholarly side and his humanitarian side.”
Page joins three other UC-affiliated entomologists in the ESA Fellows Class of 2012: Joseph Morse, professor of entomology at UC Riverside; Henry H. Hagedorn, who received his doctorate in entomology from UC Davis in 1970; and R. Michael Roe, who trained at the UC Davis Department of Entomology from 1981 to 1984 as a National Institutes of Health fellow in cellular and molecular biology.
And interestingly enough, Page is following in the footsteps of Harry H. Laidlaw Jr., who himself was elected an ESA fellow in 1991.
With the addition of Page, the UC Davis Department of Entomology now has 16 fellows who are current or former faculty members. The first was Richard M. Bohart (1913-2007), for whom the Bohart Museum of Entomology is named. He received the honor in 1947. Fifteen others followed: Donald McLean, elected in 1990; Harry H. Laidlaw Jr. (1907-2003), 1991; John Edman, 1994; Robert Washino, 1996; Bruce Eldridge, 2001; William Reisen, 2003; Harry Kaya, 2007; Michael Parrella and Frank Zalom, 2008; Walter Leal, 2009; Bruce Hammock and Thomas Scott, 2010; James R. Carey and Diane Ullman, 2011, and now Robert E. Page Jr., 2012.
Bee breeder-geneticist Kim Fondrk of UC Davis manages the Robert Page specialized genetic stock. These bee hives were in a Dixon almond orchard. (Photo by Kathy Keatley Garvey)
Springing Into Action at the Laidlaw Facility
It's not spring, but don't tell that to the folks at the Harry H. Laidlaw Jr. Honey Bee Research Facility at the University of California, Davis.
Today bee breeder-geneticist Michael "Kim" Fondrk mowed the lush green grass around the apiary. Bee breeder-geneticist Susan Cobey, manager of the Laidlaw facility, continued preparing for a series of specialized bee courses. She tended the hives with beekeeper and junior specialist Elizabeth Frost and beekeeper Tylan Selby, a first-year entomology student.
The conference room buzzed with ideas for an upcoming tour. Following the meeting, Extension Apiculturist Eric Mussen of the UC Davis Department of Entomology faculty talked "bees 'n almonds" with Tabatha Yang, education and outreach coordinator for the Bohart Museum of Entomology on the UC Davis campus. Yang will be conducting the first of what will probably be many tours at the Laidlaw facility.
The tours will include the half-acre Häagen-Dazs Honey Bee Haven and the quarter-acre Campus Buzzway. Both are bee friendly gardens that will provide a year-around food source for the bees, and educational opportunities for visitors.
Meanwhile, oblivious to the people activity, the bees continued to gather pollen and nectar from nearby almond trees.
"The queen bees are really busy laying eggs," Cobey said. That means more workers, more drones and more queen bees.
During the peak season, a queen bee can lay 2000 eggs a day, Cobey said.
That's definitely springing into action.
Kim Fondrk
Examining Almond Blossoms
Lovely almond blossoms
Italian Bee
White Gold
Almond pollination season is approaching, and with it, come concerns.
"A concern each year...is how many honey bee visitation hours occurred during bloom," writes Extension Apiculturist Eric Mussen in his newly published edition of from the UC Apiaries.
Mussen, a former New Englander who has seen dozens of almond pollination seasons in California (he's been a member of the UC Davis Department of Entomology since 1976), says California now has approximately 710,000 acres of almonds. Each acre requires two hives for pollination.
Since California doesn't have that many bees, beekeepers from around the country truck in their colonies. The going rate per hive over the last several years has ranged from $100 to $150.
Generally, California's almond pollination season starts around Feb. 10, Mussen says, and ends around March 10. That takes into account the early, mid- or late varieties that bloom at different times. However, the pollination period for each individual orchard is around 10 days.
The flight hours of a honey bee during almond pollination season? Approximately nine hours a day over a 10-day bloom period.
And what are flight hours? Mussen defines "flight hours" as "the number of hours above 55 degrees when the wind is less than 15 miles per hour, given a sufficient level of sunlight without rainfall."
"I believe that if the tree varieties overlap well in bloom, the bees usually have moved the pollen around in the morning and early afternoon on good flight days," he writes in his newsletter. "That probably requires only about four hours a day."
Of course, poor weather can interfere significantly with "fertilization and nut set," Mussen says, "but it would not be the fault of the bees."
As a service to beekeepers and growers, a retired beekeeper posts information on the Almond Board of California Web site indicating who's renting colonies and who needs pollination.
Meanwhile, check out the images below of UC Davis bee breeder-geneticist Kim Fondrk in a Dixon, Calif. almond orchard. Fondrk manages the Honey Bee Pollen Hoarding Selection Program at the Harry H. Laidlaw Jr. Honey Bee Research Facility, UC Davis, under the direction of Robert E. Page Jr., Arizona State University. Fondrk and Page moved the bees from Arizona to California several years ago.
Almond Orchard
Kim Fondrk
Honey Bee
It All Bee-Gan at UC Davis
The "honey bee reproductive ground plan" hypothesis that originated two dec
Page, emeritus professor of entomology at UC Davis and now founding director of the School of Life Sciences at Arizona State University, and his collaborator Gro Amdam, are featured in the Oct. 23rd edition of Science, a publication of the American Association for the Advancement of Science.
Writing in the behavior ecology section in an article headlined, "Sex and Social Structure," journalist Elizabeth Pennisi related that the scientists' research "has shown that reproductive traits help shape a honey bee worker's role in life and that ovaries are active players in the process-even if they play little role in reproduction in worker bees."
The specialized tasks "have their basis in what Amdam and Page call a reproductive ground plan," she wrote. Their work has provided a framework and tools to study division of labor, which now "converges on two genes that may explain both ovary size and behavior."
Page and Amdam, an associate professor in the School of Life Sciences and Norwegian University of Life Sciences, believe that genes and hormones likely control social roles as well as longevity.
Their research centers on the role of the ovary in honey bee colonies, and how the worker bees partition the labor of the colony with duties that include rearing young bees, constructing the nest, foraging for pollen and nectar, and processing the food.
Page, a pioneer in the field of evolutionary genetics and social behavior of bees, has long marveled at how highly social bees are. Worker bees, or infertile females, instinctively divide up their roles to run the hive, freeing the queen to lay eggs.
The worker bees serve as nurse maids, nannies, royal attendants, architects, builders, foragers, guards and undertakers.
But why are some colonies high-pollen collectors and hoarders, while others aren't?
His research on high and low pollen hoarding strains that began two decades led to the "reproductive ground plan" hypothesis. Page continues to keep his specialized bee stock, managed by bee breeder-geneticist M. Kim Fondrk, at UC Davis.
This is exciting research.
As Page told us: "The reproductive ground plan research is integrating developmental biology into insect sociobiology. It is completing the synthesis by looking for the signatures of levels of selection above the organism, at the level of the genes, physiology, and embryogenesis. It is substantiating the superorganism."
UC Davis is the hub for the development and maintenance of the high and low pollen hoarding strains of bees "that have been fundamental in testing the reproductive ground plan hypothesis and understanding how selection on colonies affects different levels of biological organization from genes to societies," he said.
Page, who received his doctorate in entomology from UC Davis, retired from UC Davis in 2004 to develop the School of Life Sciences in the College of Liberal Arts and Sciences at Arizona State University.
Page and Amdam are the co-principal investigators on a federally funded project directed by UC Davis entomology professor James R. Carey. Carey directs the Biodemographic Determinants of Lifespan, a National Institutes of Health/National Institute of Aging-funded program involving scientists from UC Davis, UC Santa Barbara, UC Berkeley, Stanford and seven other academic institutions in the United States, the United Kingdom, and Greece.
See more information on the UC Davis Entomology Web site.
Hives of International Interest
Queen Bee
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