May 26, 2025

Honey Bee Genetics & Breeding with Dr. Robert E. Page (335)

What do honey bee genetics mean for the everyday beekeeper—and how do researchers untangle traits like foraging behavior, hygienic tendencies, and even pollen hoarding? In this episode, Jeff and Becky sit down with Dr. Rob Page, one of the world’s leading experts in honey bee genetics and author of the newly updated Honey Bee Genetics and The Art of the Bee.

Dr. Robert E. Page, JrWhat do honey bee genetics mean for the everyday beekeeper—and how do researchers untangle traits like foraging behavior, hygienic tendencies, and even pollen hoarding? In this episode, Jeff and Becky sit down with Dr. Rob Page, one of the world’s leading experts in honey bee genetics and author of the newly updated Honey Bee Genetics and The Art of the Bee. Rob shares stories from his early influences, including his time with Harry Laidlaw and the legacy of Walter Rothenbuhler, and how his fascination with bee behavior grew into a groundbreaking career in genetic research.

They also discuss Rob’s engaging new textbook, why he pivoted away from queen rearing content, and the importance of mentorship and collaboration in science. For beekeepers interested in breeding better bees, Rob emphasizes the power of careful observation, good recordkeeping, and realistic expectations.

Listeners will also hear about Rob’s video lecture series based on his book The Art of the Bee, created to make bee biology and genetics accessible to students and enthusiasts alike. Whether you’re raising local queens or just curious about what makes a colony tick, this conversation is a rare chance to learn from a pioneer whose research continues to shape modern beekeeping.

Websites from the episode and others we recommend:

 

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335 - Honey Bee Genetics & Breeding with Dr. Robert E. Page

 

Ken Coyle: Hello, it's Ken Coyle, and I would like to welcome you to today's Beekeeping Today podcast. I'm a judge at the Royal Agricultural Winter Fair, one of Canada's premier hunting shows. Hope to see you there sometime soon.

Jeff Ott: Welcome to Beekeeping Today podcast, presented by Betterbee, your source for beekeeping news, information, and entertainment. I'm Jeff Ott.

Becky Masterman: And I'm Becky Masterman.

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Jeff: Hey, a quick shout-out to Betterbee and all of our sponsors, whose support allows us to bring you this podcast each week without resorting to a fee-based subscription. We don't want that, and we know you don't either. Be sure to check out all of our content on the website. There, you can read up on all of our guests, read our blog on the various aspects and observations about beekeeping, search for, download, and listen to over 300 past episodes, read episode transcripts, leave comments and feedback on each episode, and check on podcast specials from our sponsors. You can find it all at www.beekeepingtoday.com. Hey Ken Coyle, thank you for that great opening from the North American Honeybee Expo. Boy, that seems like a long time ago now, Becky.

Becky: I thought the opening was from Canada.

[laughs]

Jeff: I was checking my notes, and Ken, I apologize, if you want to tell us where in Canada you are from, I'll make sure you get tagged properly on our listener opener notes or website.

Becky: Let's just consider it all of Canada has now opened the podcast today. That feels good.

Jeff: I wish, but that would be against the rules of the--

Becky: The orange crayon?

Jeff: -listener opener page. Yes, the orange crayon. Becky, it's May. Man, oh, man, I can't believe it's May already. Last time we talked, you were getting set to do some divides.

Becky: Oh, I've been dividing for a while now.

Jeff: Have you?

Becky: Yes, it's been very, very exciting because I was asked-- A year ago, I talked to Brainerd Minnesota Beekeepers Association and I talked to them about modifying the Demaree method for divides because that's actually an anti-divide method, and they're like, "Come back and talk to us about it." I did a deep dive on divides, and I am having so much fun exploring all the different ways that you can divide a colony. Our regional beekeeper, Ang Roell, had a SARE Grant about runaway splits, and modifying the Demaree method is something that Meghan Milbrath of MSU does regularly. Anyway, so I'm in the middle of-- not in the middle. It's like a longest divide season ever. What about you and your divides?

Jeff: I call them splits, but that's all right.

Becky: Oh, that's a problem.

[laughter]

Jeff: We'll start there. I don't have any specific way. I haven't settled on anything, and I'm really curious. I think you and I should do an episode on the Demaree or Demaree method. I think that sounds really interesting, but most of mine have been basically walkaways and make sure pull frames, and then usually with a purchase queen and put the purchase queen in the nuc or the small lower box and call it good.

Becky: Wait, a walkaway without raising the queen?

Jeff: Correct. A walkaway with a purchase queen.

Becky: So not really a walkaway then. It's a walkaway--

Jeff: It's a skip away.

Becky: It's a skip away. It's check after a few days away and figure out which one had the queen?

Jeff: I'm walking a tight rope here. I carefully inspect a frame and make sure the queen's not on it or the brood and everything else, with mostly cut brood. I like separating into nucs and then a couple of frames of food and pollen, and then a couple of empty frames of drawn comb, and calling it good. So far, I've been good in not having a queen in that box and being able to introduce a purchase queen, a mated queen, to that nuc. That's been successful so far, although I know I should make sure that I could split my hives with the queen excluders and make sure I'm only pulling the proper frames.

Becky: No, you don't need to do that, but a lot of times when people call a walk away, they literally walk away for a month, and then they come back and they see if a queen cooked or not. Cooked and successfully made it. Everything's different. All the terminology's different, but walkaways, in my neck of the woods, would be letting those queens raise their own sister queens.

Jeff: You just take the frames, put them in a box, walk away for a month, and make sure that they both still have a laying queen, whether it was raised or the original?

Becky: That's one way to do it, but the goal with a divide is that you'll manage the one with a queen, and if you do a walk away, you leave them alone so you're not interrupting the process, so you do have to know which one has the queen. I did do an awful lot of walkaways, but I did a lot of combining at the end of last year because not all of my queen matings were great. I ended up realizing that I have to invest in really good queen stock if I want to maintain a certain number of colonies, but if you do a walkaway, usually, you know, hopefully that you're doing the walkaway in that colony, and you put that queen in a specific spot.

I was saying that Ang from They Keep Bees, their way of doing it is to actually move the whole colony to a different spot in the apiary and bring in a couple of frames, one of resources and one of brood, and that new colony is going to get all of the returning forages and the colony, they move to a different location will get the queen and then just keep going, but that new colony in that great forager rich location will get to raise their own queen. They've gotten good success in it being able to raise well-mated queens. I've noticed that if I do walkaways, which I did a lot of last year, it was really apiary dependent, which makes sense because I don't have the resources to control drones. Anyway.

Jeff: I was going to suggest that a lot of it depends on the time of year you do it and whether there's a lot of mature drones out and about flying around, and that's why there is a multitude of ways of doing your splits or your divides. That's part of the fun of beekeeping. You get to find one that works for you in your location.

Becky: Can I tell you one more way that I'm really curious about?

Jeff: Sure, I think.

[laughter]

Becky: This is in the 1975 edition of the Hive and the Honeybee, but they talk about getting into your season a little bit so there's a nectar flow, and identifying the colonies that aren't producing honey and dividing those up into nucs and requeening them. I thought that was just so interesting. The problem is, of course, if there's a disease or a pest issue that's causing your colonies to not make lots of surplus honey, that could be a problem.

If you know that you're controlling varroa and you don't see any brood issues, I think that's fascinating. Basically, it's taking your non-performers and increasing your operation by dividing them up because they weren't making honey for you anyway. We always have those colonies in the yard where we're like, "Good job. Are you going to get anything done here, or is this it?" [laughs] I don't know, I'm really tempted to do that to divide a non-performer this season, see how that goes.

Jeff: The little hive that could. I like the concept. I'm going to keep that in mind. I'm going to play with that, and if I have an opportunity to use that this summer, and I probably will.

Becky: See, aren't you glad I told you?

Jeff: Yes, I am, and I hope our listeners find that useful, too. Hey, Becky, today we're talking about honeybee genetics. It's a good time of year to talk about genetics with the father of honeybee genetics, not the father.

Becky: Exactly.

Jeff: He is the leading expert today on honeybee genetics, and he's written a lot about it and done a lot of research. Dr. Rob Page. He's out in the grain room, we'll invite him on and we'll talk to him about honeybee genetics.

Becky: Very much looking forward to it. I took a quick look at his CV because Rob Page is my graduate student days. He was very prolific. I looked, and by the time I graduated in early 2000, he had published over 120 papers.

[laughter]

Jeff: He's well published.

Becky: They're great papers.

Jeff: Oh, yes. Oh my gosh. We'll be talking to him soon about his latest update in his books and everything else. Standby right after these words from our sponsors.

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Jeff: Hey, welcome back, everybody. Sitting around this great, big, virtual Beekeeping Today Podcast table is Dr. Rob Page. Rob is one of the leading experts on honeybee genetics and behavior. He spent decades studying how traits like foraging, hygiene, and social organization are passed down through generations of bees. He's also the author of two great books, The Art of the Bee and the newly revised Honeybee Genetics.We're really glad to have Rob back to help us connect with the science of bee genetics and how this matters in our hives. Rob, welcome back to the show.

Dr. Robert E. Page: Thank you.

Becky: Thanks for being here, Rob. This is exciting. We get to talk to you for-- We've got you for almost a whole hour. We're going to learn stuff, Jeff.

Jeff: [laughs] Yes. Rob will have the opportunity to try to teach me, and you'll probably catch on more than-- No. Thanks, Rob. I really appreciate you being on the show. You really have done a lot of work, and it's really exciting to have you back. Before we really take a dive into bee genetics, can you tell us a little bit about yourself? How did you get started with bees, and how did you get settled in with this for your area of study?

Robert: I won't go back to when I was born.

[laughter]

Becky: Oh, come on. Was it in an apiary? [laughs]

Robert: I'll go back to before most of the people listening this were born. I got out of the army in 1972. I had been in for four years. I went back to school. I went to San José State University. I was a biology major, and then I took an entomology course from a very inspiring professor, I'll never forget him, Gordon Edwards. Ron Stecker, he was another one who taught. I took these as elective courses, and I really loved them. I ended up changing my major from biology to entomology. I really, really enjoyed it.

I took a course in apiculture from Ron Stecker. He had two hives on the roof of the biology building there. We couldn't do a whole lot, but I was fascinated by bees. I was coming up on time to graduate, and I still had time left on my GI bill. I didn't want to give up my GI bill, and so I decided, "But why not go to graduate school?" There's no planning any of this. This is all serendipity. Why not just go to graduate school? I applied to two, Berkeley and UC Davis. I knew nothing about either of them. I just got to the end of my undergraduate program and needed to spend some more money.

I went to UC Davis. I was going to be an agricultural entomologist type. My major professor, that's what he did, he was a good man, Oscar Bacon. He was a very, very well-known entomologist. Then I took a course, social biology and behavior, from Robert Metcalf. He was about two years older than I was. He had graduated from Harvard, he had his PhD, got a job at UC Davis. His father was one of the most famous entomologists; his name was also Robert Metcalf, and he wrote one of the textbooks in Entomology. I took this course from Bob, and I got really excited about social behavior.

At the same time, I was interested in evolution. I took a course in evolution from Timothy Prout. He was a population geneticist. He had been a graduate student of Dobzhansky who was one of the pillars of evolutionary theory. I got really interested in evolution, and then I took a course in population genetics from a guy named Bob Allard. He was a very famous National Academy plant-- He was a plant geneticist. All my breeding thinking came from not his population genetics course, which was wonderful, but it helped me, but from reading his book on plant breeding.

I realized that plant breeding, in a lot of ways, was like bee breeding. When you just think about the pollen grains and there are gametes that fly through the air. I got really excited about all of that, this combination. Then, one day, I thought I'd check out the bee lab. I went out to the bee lab at UC Davis, and Harry Laidlaw was there. He had just retired. He was 67 years old, and they made him retire at 65; there was stupid rules back then about that. Harry was really mad because they made him retire, and he was not the guy to retire.

I met him and we started talking, and I thought, "This is really interesting." I'd learned a little bit about genetics and haplodiploidy. I said, "I'd really would like to learn more genetics," so he gave me some stuff to read. Then I started going out to the bee lab, and then finally I decided, "What I can do, I can combine all these things together. I can study social behavior of bees, and I can put in genetics. I can look at population genetics of the sex determination system." I think you see where this is going. I ended up being this collection of these people who had this major influence on me was just the moment in time taking these courses shaped me, where I wanted to go.

Then I changed my major professor to Norman Gary, he was my major professor, because he was a superb behaviorist. He was a classical kind of behaviorist. That was really important in my training, too. You have to know what bees do and how to work bees and deal with them before you can ask questions about why did they do that. Then that sort of shaped me. He took me on as a graduate student. Harry Laidlaw remained my mentor. Bob Metcalf quit the university and went back to medical school.

[laughter]

He was a brilliant guy. That was how I got into working on honeybee behavior and genetics.

Becky: I mentioned, Rob, at the beginning before you joined us, that by the time I got my PhD in early 2000, you had published over 120 papers. Many, many of those papers were part of what I read while I was a student. The thing that I like most about your story is how we're talking to other researchers who are talking about the impact that you had on their careers. That's got to be one of the best parts of your journey is that you now have all these students out there, graduated students, or postdocs, or visiting scientists who have helped you in this journey, and it just continues.

Robert: You know, Becky, I did write another book, it's called The Spirit of the Hive. This book is really about my research and how it changed over time. It was influenced by every student I had, every postdoc I had, every colleague I had; they all became a part of this story. Every single one of them changed the direction I went in. Every student I had gave me back as much as I gave them, with respect to new ideas and introducing new things to my lab and to my research program, and my way of thinking about research I was doing. Every one of them did.

That's really what this book is about. It's like a journey, the work that I did. It shows that there was continuity to it from the very beginning, but it was always shaped by people. In the book, I always talk about the people who came to my lab and did the work. It set things up to move on to another question, and another question, and another question that then got back to the what I call "the spirit of the hive." It's a metaphor for two things at the same time. The're a spirit of the hive would be like my research community, as well as in the sense of Maeterlinck's question about, "What is the spirit of the hive?"

Becky: I'm not going to ask for the answer today, but if I read that book, will I get the answer to how--

Robert: Oh, yes, you'll get the answer.

Becky: No, here's my question: how you found those pollen hoarders? Is that in Spirit of the Hive?

Robert: Oh. Yes, it is, actually. The pollen-hoarding question, I give full credit to Rick Helmick. Rick Helmick, you may know Rick, but Rick was a graduate student of Walter Rothenbuhler.

Becky: I know of Walter.

Robert: Yes, I replaced Walter at Ohio State, that was my first faculty position, because he had Parkinson's and he had to retire out early. He was a wonderful guy. Rick Helmick was his graduate student. They took on this project to see if they could breed bees that were more biased towards collecting pollen. Their approach was to select for pollen hoarding, and they did. They were successful, but their whole thing was a question of, "Gee, can we select for pollen?"

They didn't follow through, they didn't do the next steps, like does it increase the number of pollen hoarders, pollen foragers, and whatnot? When I got to Ohio State, those strains were still there. They still had the high and low pollen hoarders that Rick Helmick had selected. Rick selected it, I think, just like for three generations, and then he moved on. He got his PhD and moved on to another job. They were still there, and there was a master's student of Walter Rothenbuhleler's was there, Nick Calderone. Nick also worked with these bees, and so I came in and I thought--

Nick talked to me about it. I said, "Hey, this is great. This is a great PhD project for you." Characterize some of the behavior because that hadn't been characterized. He did some really good studies looking at division of labor and whatnot, and observation hives, and he showed that the highs and lows were different with respect to their forging behavior. He went on to a postdoc, and then ultimately, he ended up in Cornell. I had these bees sitting there. I also had Gene Robinson as a postdoc. He was my first postdoc.

Gene had come in and we were sitting around and we were thinking about, there's a big field out there that really hasn't been looked at, and honeybees could really be used to study social behavior, genetics of social behavior. I took a new job at Davis, and I did not take those bees with me, because I wanted to know-- it is an old question of Stephen J. Gould. He always talked about if you roll the tape again with respect, do you get the same thing? That was the question I was asking. We went there. Kim Fondrick was Walter Rothenbuhler's technician. Kim went with me to Davis, so we got to Davis and we said, "Let's play the tape again, and let's see if we get the same thing."

We didn't know a lot because all the behavior we really had was the behavior that Nick had done, but we knew what he got. We started it all over again, and we started from scratch. We reselected the highs and lows, we confirmed the same things that Nick had, and then we went on. From that, we learned more and more and more and added stuff on and ended up doing gene mapping and all that stuff. That's where it all came from. It came from Walter Rothenbuhler and Nick Calderone, and Rick Helmick.

Jeff:: Becky, I feel like we're like the game of two degrees of Kevin Bacon or something. We're like that with Rothenbuhler and these great minds of beekeeping talking to Rob here now. It's just like--

Becky: It's really cool, and I--

Robert: The last paper that Walter Rothenbuhler ever published--

Becky: Is your names on it?

Robert: My name is on it. I knew we were getting into some things-- Gene and I published this paper in nature. People thought this was really exciting that you can actually show the genetic basis for individual behavior of bees in a colony. Everybody thought that would all just blend together, and the environment would be all that was important, and the actual genetic variation wouldn't have any effect. One day, Walter came into my office, and I loved the guy. We'd get together frequently. He came over and he said, "Rob, I did an experiment a long time ago with the brown and Van Scoy bees."

Becky: Van Scoy bees. [laughs]

Robert: Van Scoy bees. "I had something very similar." I said, "Great, bring the data." He came over and we went through his data, and I put it all together, and I analyzed it all because he was having some Parkinson's. I wrote it all up and I said, "We're going to publish this, Walter, because you did it first, and you need to get credit for it, and he was very happy about that. We established that what I had done wasn't the first; Walter had done it.

Becky: I just want to let all the listeners who might be wondering why we're so excited about Walter Rothenbuhler, but he described hygienic behavior in a couple of papers, and more. Also, I talked to on an episode of Honey Bee Obscura with Dr. Jim Tew, who knew Walter. We had this great conversation. I never met Walter, but because I studied hygienic behavior in graduate school, I had this connection to him.

Because I cited his papers all the time, so we had this great conversation. It's so exciting to hear this, and we can include that Honey Bee Obscura episode in the show notes because it's great to learn about-- In fact, one of Walter's relatives contacted us after hearing the episode and said it was just so nice to hear that people remember his work and his work was so important.

Robert: In my new book, Harry and I covered it in the first version of it. It's still central work. If you want to think about mechanisms of resistance to diseases, and his study was beautiful, the way it showed these multifactorial resistance components, it's physiological level, the anatomical level, the behavioral level, and it still holds today as far as I know.

Jeff: I look forward to talking further about this, but we do need to take a quick break and hear from our sponsors. We'll be right back with Dr. Rob Page.

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Becky: Welcome back, everybody. Okay, Rob, we could talk about this history forever and go on and on and on, but you have a book that was just released that people are very excited about. This book has a history. Why don't we talk about that?

Robert: Harry Laidlaw, as I say, he retired two years before I got to UC Davis as a graduate student. Harry had written his queen-rearing books. He wrote them with Eckert. They wrote them together. They went back, first of them was like early 1950s or whatever. Then there was the revision to that. Then Harry got tired of paying out royalties to Eckert's family.

Becky: Oh, no.

Robert: This has been a long time.

[laughter]

Anyway, he wrote the new book, Contemporary Queen Rearing. Contemporary Queen Rearing, he came to me, and I helped him solve some problems because, again, I was interested in population genetics. He was wanting to come up with this simple model for breeding bees and avoiding the problem of homozygosity at the sex locus. I said, "I think I can do that." I knew how to program, and then I knew enough population. I did these simulations and things, and I figured out what's the minimum number of colonies you need to have to avoid serious inbreeding for 20 generations, or not-- It is inbreeding, but it's not inbreeding depression, but it's inbreeding. It's the buildup of homozygosity. Anyway. That went into his book.

Becky: Homozygosity at the sex allele leads you to diploid drones, which are not viable, just so that everybody here is with us.

Robert: Yes, and shot brood, which is the problem to the beekeepers. Then, when I was at Davis, when I came back as a faculty member, because I did all that work as a graduate student with him, and he said, "You know, Rob, I want to write a text on Queen Rearing and Bee Breeding." He said, "I want it to be something that would be valuable to bee scientists but also understandable by beekeepers and bee breeders who have an interest in the biology and in understanding the genetics." It came out in 1997. Harry was 90 years old. 90 years old, and we're working on this book.

Harry died in 2003, 96, I think he was. He sat down at his desk, he started working on all this. He had this huge collection of photographs. Harry was an avid bee photographer. He took slides, and he took black and white, lots and lots of black and white. Everything was organized. He spent a lot of time at the UC Davis entomology department dark room when he was in his 80s. In there, taking pictures of beekeepers doing this and that, and beekeeping operations, and he printed them all out in these little-- in four by five black and white pictures and cataloged them in his catalog. I have all that. I got to turn it over to someone who will archive it. I've got black and white pictures of Harry Laidlaw with Karl von Frisch at the bee lab visiting.

Becky: Oh my gosh. Holy cow.

Robert: It's amazing. He'd sit there and go through all these pictures, and he'd find the negatives, and he'd make another better copy-- this was all those black and white photos that went into the book, the QueenRearing and Bee Breeding book that was published by Wicwas Press. I did the genetics and the breeding theory, and whatnot, but 2/3 of the book was what Harry put into it. A lot of that book was about basic beekeeping kinds of methods, apicultural methods of queen rearing, and along with all the pictures that shows everything. It's a wonderful book.

When he was beginning to fail, "Rob, you got to do a revision of the book. It's out of date already." This was 2002 or so, he's telling me this, 97-- "You got to do a revision of the book. You need to revise the genetics because that's what's changed so much." The queen rearing hasn't changed much since the 1880s. It's still pretty much the same thing. It's just, okay, what are you using today for a pollen supplement or whatever, compared to what it was 20 years ago? I finally got around, I've retired, and all that kind of stuff. I started thinking I really need to do it. My wife was nagging me about it. Michelle nagged me about it, "You promised Harry, you were going to do it."

Becky: Oh, wow.

Robert: I decided, "Okay, I'm going to take it on." I think I started this project maybe three years ago, maybe a little more. I just started working on it and outlining it, and going through Harry's book. Here, you can see all my notes, and pencil notes, things I was going to change. I was working through all the queen-rearing methods. I was asking Kim Fondrick, who was retired too, but he was my technician at Ohio, and at UC Davis, and at ASU.

When I came to ASU, he came with me and was my technician here, too. I just started thinking to myself, "I can't really do this because I don't do that." Harry and I would take trips out and visit all the queen producers. We did it twice, and he took pictures of everybody, asked them all the questions, and he knew everything that was going on. I don't know what had happened since the book was written. I relied on Kim a little bit because Kim works with-- Then I finally realized, "I can't do this. I don't really have the practical know-how."

That part of it is not something I've kept up with. I was provost of the university here at ASU. I had other responsibilities. I started thinking about it, just writing it as a book on genetics, bringing all the stuff on honeybee genetics, and social genetics, and make it just a textbook on that. I spent three months in Münster, Germany, and I took all my stuff with me. I was getting out of town so I could think a little bit. One of my former postdocs invited me for three months.

I started working on it in earnest there, and then I decided I'm not going to do the applied queen rearing parts, it's going to be the genetics, and I wrote it. I sent it to Oxford University Press, the same editor that had done my Art of the Bee book, and they really liked it. They passed around, they said yes, they want to publish it. Then my wife started nagging me, Michelle, "Rob, this isn't what Harry wanted. You're going to publish this in Oxford University Press, and the people that Harry really wanted to see it and read it and get aren't going to get it."

Then I asked around a bit, and also, along the way, I had asked Sue Cobey if she thought a book like this would be a value. She said absolutely. She really encouraged me to do it, and so did Kate Ihle. They both really encouraged me to write this book. Then Michelle was telling me, "You got to do it for something that will be of value to the target audience that Harry had." The academics can read the book and can get a lot out--

In fact, my German colleague told me that he's making all of his bachelor's students read the book just to learn about the genetic. The genetics is valid. I got a hold of Larry Connor at Wicwas, and I said, "Larry, what do you think about this?" He thought it was a great idea. He encouraged me to do it, so I started reshaping it into something that would be of more value to an audience that Larry and Wicwas Press would reach instead of an audience that Oxford University Press would reach. That's where we are. There's parts in it that were in the other book, in the first book, the parts that I had written on genetics and things, but for the most part, it's new. That's how I got to the book.

Jeff: What were the big changes in the genetics that you wanted to capture in the new book from the original book?

Robert: I think that the two main things that happened that I wanted to capture were all the changes that had taken place in molecular genetics. It was only just starting in the first book. There is a section in there on QTL mapping, but QTL mapping was brand new when I started doing that. Nobody was doing it. I was only doing it because Greg Hunt came into my lab as a graduate student, and he happened to have a wife who was a postdoc in Wisconsin. She was working on trying to construct a map of the rice genome. She was a plant geneticist.

When Greg came to my lab, she got a job in a lab at Davis too. We had her just write down the couple of buildings down and she had all the new technologies. She was working in the lab that had published the first map of anything, and that was of the tomato, and it was done at Davis. She had all the marker stuff. Greg worked in that lab and learned how to do all the marker techniques that you need to do to do a map, and then started applying them to honeybees.

That was here, but then beyond that, all the other stuff that came along was not in there. I wanted to bring it up to date, and then I wanted to analyze how it's been used and look at how it can be used, and assess where it ought to go. That's one of the main things I wanted to put in. The other thing was that there had been some advances in very sophisticated quantitative genetic methods being applied to breeding in different areas of breeding for resistance, for honey production, et cetera, very sophisticated models.

I wanted to look at those and figure out, okay, it was a lot of work to do. Working with these models, did they get anything out of it? Was there any evidence that there was any real advancing? Did honey yields really improve? Most of the honey yield stuff, improvements, the way I look at it, is honey yields went up in Europe when they started controlling varroa, not when they started changing their breeding methods.

I wanted to take a critical look at that and also put that in the book. Then, any other things that had come along that I thought really had shown some value with respect to honeybee breeding for characteristics that were of importance to beekeepers. I don't spend a whole lot of time on pollen hoarding in the book. We learned a lot about honeybee genetics and behavior and whatnot, but I don't think that anybody took it to the bank.

Becky: I have to ask. There are so many groups emerging across the country, beekeeping groups, who are deciding they want to start trying to control their stock, raise local bees, and hopefully improve wintering and possibly pest and pathogen resistance. It sounds like those groups need to read your book.

Robert: I think they would be encouraged. I think it's a great idea. I think that if you are a careful observer, if you use good assays to evaluate the stocks for whatever you're wanting to select, if you're realistic in your expectations, and you're systematic in your approach, I think you're going to make progress. I think the evidence of it is everywhere. When you go look at beekeeping, different beekeepers have selected stocks for color, organization of the brood nest.

These are all things that were done locally. I saw that when I'd go out with Harry and we'd go visit different areas. You went over and you visited Adrian-- not Adrian Winner. What was his uncle's name? Anyway, the winter apiaries, their bees were black, and he really liked the blacker bees. Then you went up and you visited the parks and Homer park. He liked to have really big, fat yellow queens. That's what he had. Selection at a local level like that can work.

Ernesto Guzman and I did a big study in Mexico using open mating and in a selection program. We were trying to select bees that didn't have the Africanized honeybee characteristics in an area with Africanized honeybees, and we were able to do it. It was successful. There's four to five different explanations for why, but I think that that can work. I think you don't have to have these really sophisticated, complicated computer models that I think you just call out what you don't want and keep what you do want.

There's lots of ways of doing it. I deal with that in the book. That was really Harry's view. Harry's view was keep good records, have good assays, know what your phenotype is that you want, and then control your matings. You can control your matings by mating yards. You can control your matings by instrumental insemination. If you do that, you're going to make progress.

Jeff: Regarding genetics, are we any closer to identifying specific genes that might be able to reduce viruses or identify specific behavioral traits?

Robert: Yes and no. I'll stop right there. No, I won't. I'll get that.

[laughter]

Yes and no. I think that we are identifying genes that are involved in gene networks that are affecting certain kinds of phenotypic traits. I talk about that in my book as well. Genes don't act in isolation. In fact, maybe Gene said it first, Robinson. I don't remember, but there's the ecology of the gene and the genome, that genes aren't acting independently.

Their actions depend upon a lot of other actions of a lot of other genes and a lot of other networks. You've got all these networks of genes that tend to work together, and then they cross-talk with other networks, and that's really what you're working with. When you go in and do a selection program like I did for high and low pollen hoarding, I remodeled a large number of set genes, I'm sure.

Our ability to locate and identify specific elements of that complicated network that we change limits us only to finding things that have really big effects. Sometimes those big effects are only in the gene environment in which they're working. There is no silver bullet that we're going to find the gene for this or the gene for that, and that's going to take care of it. It doesn't work that way.

Jeff: No magic switch to turn off deformed wing virus or something.

Robert: There are known cases in entomology where there are gene-for-gene actions between plants and insects, where a single gene is affecting an insect's ability to get around the defensive system of a plant. Then the plant can get a mutation that will then offset that single gene and the insect. They get into these things that go back and forth, though they're very rare. It's very rarely that simple.

Jeff: Obviously, we could have a whole podcast devoted to honeybee genetics, and if you want to, we can talk to you about it like later.

[laughter]

Really, though, but you're involved in other things too, besides your book. You have some YouTube videos, The Art of the Bee.

Robert: I wrote the book The Art of the Bee because I wanted to write something that was different. I wrote The Spirit of the Hive, and when I got done with it, it wasn't what I really wanted. It's a nice story about how science is done, but still, the audience, beekeepers, bee enthusiasts, aren't going to go read a book about how science is done. Even though there's a lot of interesting behavior and biology and everything in it, it didn't really hit the target that I wanted.

In 1996, I got a Humboldt Prize. Germany gave it to me. I spent a year in Germany on a Humboldt Prize and read a lot about Humboldt. I got really interested in Humboldt as a person, Alexander von Humboldt. He wrote some really interesting stuff. He wrote this one book that was called Ansichten der Natur, and that's Views ofNature, but it's views of nature with double meaning. It's a view of nature, like this is my view of it, and it's also the view of nature.

It's translated into glimpses, but in this book, it was a compilation of his five years of travel throughout South America, Central America, Mexico, Cuba, and where he kind of got this composite view of how-- Basically, community ecology was something he invented. He showed the connectedness of everything. Then he would write a chapter on some interesting theme. It wasn't about any one particular-- It wasn't on biogeography of this particular plant in Chile. It wasn't like that. It would have some other kind of theme to it.

He would wrap that into every chapter and sometimes even paragraphs. He would have botany, zoology. He'd be measuring magnetic flux. He would be measuring atmospheric pressure. He even put anthropology in there because he was studying the natives and showing their influences on the environment. Then in the end of the book, he challenged other scientists. In order to get your message out to the people, paint it. He was an artist. He challenged to paint. You could paint with a paintbrush, or you could paint with a narrative. That was his challenge. There was a guy in the mid-early 19th century. His name was Edwin Church. He was an American artist. He was really impressed with Humboldt, and so he retraced in South America the track that he had taken. He saw all these things that Humboldt had seen. He came back and he painted one picture. It's huge. It's called The Heart of the Andes. Look it up. The Heart of the Andes. It's like five by nine feet. I think it's sitting in the museum of-- not modern art. What's the big one in New York?

Jeff: Metropolitan.

Robert: Metropolitan. It's sitting in there. Anyway, this is just an incredible thing, but it was everything. None of that exists. It was just him making a compilation of the ecology. There's anthropology in it. All this is in there. I got fascinated. I said, "I want to do that." [laughs] I wrote the book without an outline, but I'm not going to have an outline. I'm just going to have a theme for each chapter, and it's going to be just an interesting topic. Then I'm going to weave in everything into it.

Every chapter has something on bee behavior and anatomy and physiology, and genetics, and weeding it all into each chapter around some kind of a basic interesting theme. Then I gave a copy of it to president of my university, Michael Crow, who's an incredible man. I worked for him as his provost and dean, and everything. Anyway. He said, "Rob, and I'm already retired. You got to turn this into a course. You got to do an online course on this." I said, "Okay." I took that as a challenge now from him.

He's been very good to me, and so I was going to take it on. I sat down and I started thinking about what to do. I went over and I talked to these people at what we call the EDPlus, which runs the ASU online. They got really excited about it after Michael told them that they would give me everything that I wanted to do it. [laughs] I had all the studio time in the world. I had all these people working with me. I then went and I put together 50 videos. They vary from 10 minutes to 30 minutes, or 8 minutes to 30 or something.

I did it then to complement the book. It makes the book even more accessible if you do the videos, and that will help students. It'll direct them into what I thought was interesting. I teach this course. I have 100 students at each semester that take it online. They use the videos to complement the book. It was fun. It was a lot of work to put all those videos together and go into the studios. If you look at it, my hair's long sometimes and it's really short sometimes, but I always worry they're a black shirt or a black sweater.

Jeff: They are nicely done. They are easy to watch.

Becky: They're beautifully done.

Jeff: Yes, they are.

Becky: In this course, have you had students who maybe got the honeybee bug or the entomology bug because of what they've learned?

Robert: I've had some tell me that they want to learn more and do more. They've asked me, "How do I get bees?" and things like that. This is an online course. When I have my office hours online, I say, "Where are you from?" "I'm in New York right now." "Oh, I'm in North Carolina in Asheville." The hurricane went through, and she hadn't had water in her house in four weeks. One in New York City was working at Starbucks, and she was watching my office hours at Starbucks in the store room online.

Becky: Oh, my gosh.

Robert: They're all over. They come in from everywhere. It's really interesting.

Jeff: Interesting.

Robert: Columbus, Ohio, Madison, Wisconsin. Several from Texas. These are the few that actually come to my office hours. I get like five or six each week.

Becky: Can we include a link for the next registration of that course?

Jeff: We'll include it in the show notes. Yes, absolutely.

Robert: They can take it off ASU online. It's offered.

Jeff: Rob, it's been a true pleasure having you back on the show to talk to us about honeybee genetics and everything that goes into it. I was afraid that you would overwhelm me with the deep dives into specific genetic terminology and theory and realities, but I'm glad that you didn't. I am inspired to continue my exploration of honeybee genetics, just by talking to you. Great having you on the show.

Robert: Thank you.

Jeff: You're welcome.

Becky: What a pleasure. Thanks, Rob.

Robert: Thank you.

Jeff: I can see how much he really enjoys his area of study, the genetics, and how it all works together. That's a fantastic discussion.

Becky: Yes, that was so much fun for me to be able to talk to him and learn from him because he honestly takes his research in a very good way, so personally. He's created this network of not just amazing data and results, but also relationships. That was just a fun road to travel down.

Jeff: That about wraps it up for this episode. Before we go, I want to encourage our listeners to follow us and rate us five stars on Apple Podcast or wherever you download and stream the show. Even better, write a review and let other beekeepers looking for a new podcast know what you like. You can get there directly from our website by clicking on the reviews tab along the top of any webpage.

We want to thank Betterbee and our regular, longtime sponsors, Global Patties, Strong Microbials, and Northern Bee Books, for their generous support. Finally, and most importantly, we want to thank you, the Beekeeping Today Podcast listener, for joining us on this show. Feel free to leave us questions and comments on our website. We'd love to hear from you. Thanks a lot, everybody.

[00:57:15] [END OF AUDIO]

Robert E. Page Profile Photo

Robert E. Page

PhD, Author, Professor Emeritus, Geneticist

Professor Emeritus Robert Page joined Arizona State University in 2004, after retiring as Professor and Chair Emeritus at the University of California Davis, 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 is outlined in his publications:
- “Queen Rearing and Bee Breeding” (1997, with Harry H. Laidlaw),
- "The Spirit of the Hive,” Harvard University Press (2013) and
- "The Art of the Bee", Oxford University Press (2020)

His 230+ research papers have been cited more than 20,000+ times. He is an elected member of the American Academy of Arts and Sciences, the Brazilian Academy of Science, Leopoldina - the German National Academy of Science, and the California Academy of Science. He was awarded the Humboldt Research Prize, is an elected Fellow of the Entomological Society of America, and recipient of the both the Eastern and Western Apicultural Society Research Awards.

Professor Page retired in 2019.