Secrets of the Honeybee

Ask A Biologist Podcast, Vol 131
Podcast Interview with Robert Page
Robert Page at the ASU Bee lab looking at a honeybee swarm.

Dr. Biology:


This is Ask A Biologist, a program about the living world, and I'm Dr Biology. In this episode, we're going to talk with a biologist who has spent more than 40 years studying one animal, an animal that has six legs, flies and makes something sweet that most of us love to eat. By now you probably know that I'm talking about the honeybee. What you may not know is, in those 40 years of research, our guest biologist has likely worked with more than a half a billion honeybees. That would be honeybee workers, which are all female. There are also the hundreds of queens and male drones that have been part of the research. 


So, what has our biologist learned in their 40 years? That is what I hope we'll explore in this show with our special guest, Robert Page, an award-winning entomologist. During his career, he has published over 250 research papers that have been cited, which means they've been included as references in over 18,000 other scientific publications, and, to add some more writing to his resume, he's also published several books about his favorite insect. Now I suspect we will learn a lot about honeybees that we didn't know and a bit about a career in science that has spanned four decades. Welcome to Ask a Biologist, Rob. 



Well, thank you, I'm happy to be here, Dr. Biology. 

Dr. Biology:


So, when we talk about honeybees, it's one of those things that there are some people that have an aversion to bees, but in general, people love in particular honeybees, because honeybees are only one type of bee that's out there. There's a lot of different kinds of bees. 




Dr. Biology:


What is it over these 40 years? You've been doing this for 40 years. What is it that has surprised you the most about the honeybee? 



I think the thing that has surprised me the most about the honeybee is that not only have I been studying honeybees for over 40 years, I've been studying one thing about honeybees for over 40 years. My focus has been on how they can evolve to have such a complicated society. Their social system is very similar to ours in a lot of ways, and how did they evolve this social system where there's no one in charge? The queen isn't in charge. The queen bee basically just lays eggs. 


There's no president of the society or any true king or queen in the sense that we think about kings and queens. Every single individual makes a decision independently of others. Independently in the sense that there's no one who tells anyone what to do. They all decide on the basis of what they themselves can perceive sense in the nest, what the environment is, and then they respond to it. And what you get from that is this incredibly organized, coordinated society. 


So, I have studied it from that level, looking at the different behavior, looking at how individuals interact with each other and form this society and build these wonderful combs that they build. But also I've been looking at the genetics. There has to be genetic basis for the behavior. What are the genes that are involved in the behavior that glues together this social group, and then how did it change over time as these very complicated societies evolved? So my biggest surprise has been that every time I've answered a question, I've had four brand new ones arise. So I look at where I am today and I have far more questions that are left unanswered than I had in the beginning, when I started doing this research. 

Dr. Biology:


Right, it's kind of where you start peeling back the layers of the onion right. There's always more going on. 



But the big difference there is that when you peel the layers of an onion it stinks. When you peel the layers of honeybee, behavior, it's just a fascinating, incredible adventure that you start on Right and sweet, and sweet. 

Dr. Biology:


When you talk about these really amazing societies that the honeybees have, it's female-dominated, which is another thing that a lot of people don't think about. So you have male drones, but they're not around long, are they? 



I would not say that it's female-dominated. I would say that the males and the females have different kinds of things that they do and this has all been part of the social structure, the reproduction that's needed in order to maintain the societies and the lineages. So, there's two kinds of females. There are females that are the worker bees. They have stingers. They go out and forage and they collect pollen on their legs and collect nectar from the flowers that they turn into honey, and they're responsible for the vast majority of the tasks that we observe, the way that they can control the temperature of the nest. They can collect nectar, cure honey, build combs, they do all those things. The other kind of female is a queen, and typically there's one queen in a honeybee colony and anywhere from 10 to maybe 40,000 workers. No one is in charge, no one dominates anything, the workers. They build the comb, they feed the queen protein we call it royal jelly which then she turns that into eggs and she lays eggs and cells only after the workers have prepared the cells. If they don't prepare the cells, she won't lay them and thereby they control her egg laying by how much they feed her and preparing the cells for her to lay in. So she's not in charge of anything. 


The males, their task in life is to leave the nest, the hive, every day in the early afternoon and go, fly through the air and they locate places where virgin queens, as queens that have not yet mated. They fly there and they encounter males and the males mate with them, which is pretty spectacular. My understanding the mating is an incredible. I call it the song of the Queen. So she will mate with, you know, up to 20 males as she flies through the air, and each of the males, when they mate with her, in the act of mating, they become paralyzed and they fall to the ground and die. 


And so for them, that's the one thing that they're designed to do, and they're designed very well to do it, because they have very special features. They have large eyes that have more of the individual eyes. If you look at an insect eye, you'll see that it has lots of smaller eyes that are called eye facets but they have more than the workers, which gives them a better ability to detect movement when they're flying through the air, so they can locate Queens. And they also have antennae that have Receptors for sensing the odors of the Queen. The Queen produces these odors called pheromones, so they fly through the air and they're just really designed just to fly through the air and detect a queen and mate with her. 

Dr. Biology:


So they have a very important function, but it's not the same function that the workers have, and the Queen's function is not the same as the workers you know I did some reading and I think I have this right that the average honeybee colony has as many neurons as a human brain, and If that's true, that seems like a lot of brain power. So how are Honeybees using their collective brain different from humans? 



Well, that's an interesting question and I think you're pretty close to being right with respect to the number of total neurons in a hive and I think about those things sometimes. I think about what is the brain of an insect colony. What does it have? If you take each and every single one of those, let's just say, 40,000 workers in a colony, each one of them has a very different set of experiences. Every one of them has a memory, and they do have memory and they do learn. We can test it. We have ways of knowing what they learn. We have ways of teaching them things and then asking them questions back about what they learned. So, we know they learn. Every one of them has learned something different. 


So, if you went inside of the nest and you were looking at the organization of the nest, the things that need to be done, you have this collected memory in there of the things that need to be taken care of and the things that have already been done, just as we have in our own brains. 


We know what we've done and we know what needs to be done, and we know what the inside of our house looks like and where to get things, and they do too, but they do it as a collective and then if you look where they forage, every individual can be foraging out Up to five miles in different directions and they come back and they perform dances that inform the other Individuals as to where these resources are located, the direction and the distance. 


They have individual memories about the landmarks that are out there, that associated with the Location of the nest, as well as the location of the resources. So, if you put that all together, you have an incredible map of the environment and they do communicate to some extent with each other, to where they can transfer that information around, very much like In our own brains how we assemble information and act on it. Colonies of honeybees can forage as a collective unit optimally, as if they have information that's global, meaning that all individuals share the same information about everything that's out there in the environment. Even though they don't, there are mechanisms that they have whereby colony can, as a foraging unit, change over the course of a day from foraging on resource flowers that are less profitable to them, has less sugar in them to bring back and shift over to forage for those that are more profitable. They can do that as a collective unit, though no one individual has all that information herself. 

Dr. Biology:


Wow, you know you mentioned the bee dance, the waggle dance, and I just want to put a little plug in that we have a really cool game called the waggle dance game on Ask A Biologist. So, if someone wants to learn how bees communicate between themselves, that's the way to get started and, quite honestly, once you've played that game enough, you Actually will understand what they're saying, which is pretty cool. Yes, so you know you talk about this "collectiveness" and one of the things I realized is an advantage of the bees is I can only go one direction and come back right. Wow, as if you have 40,000 bees now. Not all of them are going out foraging, but of all those foragers are going out. As you mentioned, they can go in all directions, and so they get this collective information much faster than if I had to go out to all those different locations as an individual. Are there other insects that do that? 



Well, of course, there are other social bees, and there are also social wasps. So in order to be doing that, you'd have to be living in a social group. 

Dr. Biology:


That's some ants. 



Yes, ants as well ants, bees and wasps and what they're doing is what we call central place foraging. They're foraging from a central place and for that to happen, you have to have a nest. So you think about you have a nest, you have Resources scattered all around the environment, all around Central place foragers then forage out from their nest, they find resources, they bring them back in, and it's a huge amount of collective information. If you were to be able to collect all the information that every individual has, again, be it an ant, be it a wasp, or a termite, termites do the same thing. They form very large social groups. 

Dr. Biology:


Hmm, all right, so you told me that one of the things that has always impressed you with the honey bees is that every time you find an answer, it seems to open up multiple new questions for you. Is there Something that you recall in that 40 years that really surprised you? I mean, because we have a tendency to think we know how certain animals will behave and what they will do, and then the really fun thing about science is find out that we were really all wrong. So is there something that comes to mind that you were just like wow, this is really, really surprised me. 



There is one thing, but I have to also say that I tell my students that science is a self-correcting process. You have hypotheses. Most of your hypotheses are demonstrated to be wrong, so you have to be willing to accept that you were wrong. And again, any scientist that tells you they were always right, nothing went wrong while they were on their path to discovery, well, they're not telling you the truth. I've been wrong much more than I've been right. But the one time that I was right that was really the most exciting part was I was studying the reproductive system of the worker honeybees. Workers have ovaries. They don't normally lay eggs. They're what we call facultatively sterile. That means they can be sterile or they're not necessarily sterile, under certain conditions, but normally they don't lay eggs but they do still have ovaries. 

And I was studying the ovaries of some bees that we had done some artificial selection for. We selected for bees that collect a lot of pollen and bees that collect a lot less pollen, and it was a selectable trait. So I had some colonies that had bees that were high pollen collectors and some colonies that had bees that were low pollen collectors. We were trying to figure out all the different things that were different between them, to try to figure out maybe a mechanism associated with forging decisions of bees to forge for pollen or nectar. And we were looking at the ovaries of workers. 

I can't remember exactly why, but I had an undergraduate working in my lab and she was a sophomore, I believe, undergraduate and she came to this other person I was working with a postdoc who was in my lab at the time and she said you know what? Those bees that collect more pollen, they have bigger ovaries. And so then, all of a sudden, we started thinking about it. We hadn't noticed that. It was her discovery that we found it. So, we started thinking could it possibly be that worker bees that have larger ovaries are also going to be more likely to be the pollen collectors of a colony? And we subsequently did studies and found out that that was the case. What we called reproductive status of an individual and their forging behavior, and you could think about it in the context of how behavior changes with reproductive cycles and reproductive state. 


The one I like to use is look at a mosquito. Mosquitoes, when they first emerge as adults, when they first come out of the water and they emerge as an adult, the first thing they do is they fly around and they find flowers and they take in nectar. They suck nectar because they need carbohydrate, they need the sugar resources to give them energy. Their ovaries are not yet developed, they're still in a developing stage. After they feed on sugar, then their ovaries start maturing. 


They start maturing, they start developing, they get bigger and then their behavior changes. Instead of flying around searching for flowers and nectar, they start flying around searching for blood meals. Like you and me, they become sensitive to carbon dioxide, they become sensitive to body temperature and then they will alight on you and they will suck up a blood meal. They'll fill up with blood. Their stomach fills up with blood After that. Now they're full, they're satiated, their ovaries are still enlarged. They're ready now to make eggs. 


Then their behavior changes again. Now they start looking for a cool, dark place on a vertical substrate where they will then just sit and make eggs. In your own house, one of the places they like to sit is just right behind your toilet, in your bathroom, because it's humid, it's dark, they're full, they process, they make eggs, then, after the eggs are fully developed, now they're getting signals that change their behavior again. Now they fly around looking for surfaces of water to lay their eggs on. Once they lay their eggs, their behavior changes again. Now their ovaries are shrunken again, and now they become nectar foragers again because they need to get more sugar in order to start the cycle again. So, basically, what we found was that pollen foragers are like the blood meals seeking mosquitoes with larger ovaries, and the nectar foragers are like those with the smaller ovaries that are searching for carbohydrates. 

Dr. Biology:


You know, I wasn't even thinking about that, and of course there are other animals out there that have a similar cycle. 



Yet, like humans and turtles, it's actually called a gonotropic cycle, gonotropic having to do with reproductive organs. 

Dr. Biology:


Now as a scientist, you've been an educator. You've taught. You're also a writer. You've written several books. One is the Spirit of the Hive and the other one is the Art of the Bee. There are others you have, but these are the most recent ones. If someone wanted to learn about the honeybee and they had to pick one of those books, which one would you recommend? 



I'd recommend the Art of the Bee. 

Dr. Biology:


Okay, and why? 



Well, I wrote the Art of the Bee to be a basic honeybee biology textbook, but not in a traditional textbook. Traditional textbooks present you with information that is in what I call organizational chunks, so you might learn about taxonomy of bees. To start with, that would just be being able to identify bees. Then you might learn something about their biogeography, that is, where they're distributed around the world and what do you find them in tropical areas or on islands, so you'd study their biogeographical distribution. Then you might start studying something about their anatomy, about their physiology, and then you maybe learned a little bit about their neurobiology and then their behavior. Then, at the end, you'd learn something about how all this stuff leads to social behavior. I wanted to have a book that was focused more on interesting questions about bees and that then brought in all of those different levels of organization into the same chapters, into the same paragraphs, instead of presenting it in layers, present it together, kind of as a case-solving system where you get everything and you solve a problem using all the different tools. 

Dr. Biology:


Right, kind of like just-in-time learning right? You have a question and you need to find the answer for that particular question, and it might require a few different pieces to solve it. Yes, out of the millions of bees that you've observed, have you ever had any that were somewhat unique? I mean the workers that just behaved differently than others, that actually stood out. 



As individuals, not their behavior as colonies. Yes, I had one case where I had a colony of bees that we give them combs the frames where they've constructed the comb. We'd give them a comb and they would chew all the wax up and spit it out the front of the entrance. You'd walk around and you'd see piles of wax sitting out front where they would strip all of it down to what we call the mid-rib of the comb, the very center part of the comb. The comb's constructed on both sides and it goes out in two directions, but in the very base of those two sides there's what we call the mid-rib and they'd strip everything down to the mid-rib. Then I found out that there were a couple of other colonies in the apiary doing the same thing. So then I went and I'd look back at our mating records and breeding records and found out that the queens in those were sisters of each other. So that told me hey, you know, we probably have a genetic trait here for destroying this comb. 


It just popped up in a breeding experiment we were doing and so I told my technician at the time. I said I want to save that. That's a really interesting mutation. So let's try to save it, but we couldn't. How are you going to save something that breaks its comb down? There was a pathological trait that popped up. There was a mutation somewhere along the way that affected their sensory system, probably to where they over-responded. They normally do respond to certain signals and tear wax down, but they had an over-response to it to where they just tore everything down and we lost it. We called it a line of bees because it was one family that did this, but that was the most striking single thing. 

Dr. Biology:


Right, and so by tearing down all that wax and everything they basically they had no way to raise a new colony. 



They could not raise new bees because they wouldn't accept any comb to raise them. 

Dr. Biology:


Wow, self-destructing colonies.



It was definitely pathological.

Dr. Biology:


Wow, that is an interesting one Now, rob on, ask a Biologist. My scientists don't get to leave before I ask them three questions. The same three questions. So are you ready? 




Dr. Biology:


The first question is when did you first know you wanted to be a scientist? 



I wanted to be a biologist since I can remember. I believe when I was about five or six years old I wanted to be a biologist. I organized a little biology club in my neighborhood. One of the children on my block had a little clubhouse that his parents had built him and so we turned it into a biological laboratory. So that was what I wanted to do since I can remember. But I didn't really get into working with honeybees until about 1973. I took a course when I was in college and I was a biology major. I took a course in apiculture just as an elective course and that then really excited me. It excited me about bees and it excited me about insects. So, I took other insect courses that were being offered and every insect course I took I liked and I took another and another and before I knew it I had all the courses I needed to graduate with a bachelor's degree in entomology. 

Dr. Biology:


Ah yes, the study of insects. Okay, Not to be confused with etymology. 



That's correct which is the study of words. 

Dr. Biology:


Right, okay, so 1973. Did you go just straight from high school into college? 



No, I graduated from high school, and I went away to college in Los Angeles for one year, and then, when that year was up, there were circumstances where I could not go back for another year, and so I worked, and this was during the Vietnam War. So, I was working and I lost what they called a student deferment. Back when we had a draft which only drafted males at the time, men. You had to either have a deferment or you were classified as one a one a meant that they could draft you into the army at any time. A student had what was called a two s deferment. I lost my two s deferment because it was no longer in school, and I became a one a and I ended up in the army. 


When I went into the army, since I still had an interest in biology, I wanted to go into what they call the medical service corps, so that's what I asked for, and they sent me to be trained as a combat medic. That wasn't exactly what I had in mind, being a combat medic, and so I then decided that I would go to officer candidate school. So, I put in the paperwork and they accepted me into officers candidate school. I was only 19 years old at the time and I said that I wanted to get a commission in the army, in the medical service corps, which they also had. I did not get that as an enlisted man, I thought maybe as an officer. So that was my first choice and my second choice was engineering. 


They have an engineering branch of the army. That is very important, and my third choice was the signal corps, that's for communications. They sent me to infantry, where I stayed in the army and additional three years. So, I spent four years all together, and the last three years I was stationed in Germany in an infantry unit. I got out of the army in late 1972. And that was when the Vietnam War was starting to come to an end, and so they let people get out more easily and earlier, and so I asked for that and I went back to the university. I went to San Jose State University and I picked up my biology interests and had a biology degree program that I then changed to entomology. 

Dr. Biology:


All right, you have this passion for insects in general. When was it that you figured out that honeybees were your favorite insect? I mean, did they speak to you? 



Well, they talk to you. You can listen to them if you understand their dance language. It's almost like Dr. Doolittle, he could talk to them as well, but you can't talk to the bees, but you can listen to them. But my interest focused on bees shortly after I went to graduate school. 


When I finished my bachelor's degree in entomology, I still had money left on my GI bill. The GI bill was something that was really beneficial for being in the armed services, because when you got out they gave you three full years of paying all of your college costs. So I still had some time left on my GI bill and I went to graduate school at University of California Davis and while I was there I found out that they had a program in apiculture that is, the culturing of bees, and so I took another course and I decided that that's really what I wanted to work on, and I had some wonderful mentors who helped me along. I had three professors who took an interest in me, because I took an interest in them and they really helped guide me and direct me to establish a research project that involved honeybee behavior and genetics, and that's what I've done ever since. 

Dr. Biology:


The second question is where I'm a little bit evil because I take it all away. Because we know, for 40 years you've been passionate about this, you've been doing it for a very, very long time, but in this thought experiment, I'm taking that away. I'm taking away your teaching and taking away your writing. And if you could do anything else, if those things aren't something you're going to be able to do, what would you do or what would you be? 



The only other thing I ever wanted to be something that you, Dr. Biology are superb at and I'm not. I wanted to be a photographer. That's where I really would have gone and I toyed with it. At one point in time, when I got out of the army, I was actually thinking about going and learning more about photography and trying to make a career of it. That's really my other passion. 


But I have to say that over the last couple of weeks, I have been looking at thousands of pictures that I have taken over the years. It's something that you do when you get older. You find out that you collected all these things. You never looked at them. You took the pictures, you had them printed. You threw them in a drawer. Well, I have bins full of them that I now want to start cutting down on how much storage I'm using. So I have been going through these pictures and the one thing that I have discovered is I made the right choice. Going into science, going into biology, was the right way to go, because I have a lot of really bad pictures that I've taken over a lot of years, and I look at you and you're laughing, and you are a superb photographer. I am very impressed with the work that you do for Ask a Biologist, as well as your other things that you're engaged in. 

Dr. Biology:


Well, I thank you. I think you're my first guest that has actually picked something that I actually do in my life. I actually do a lot of, and I appreciate the fact that you enjoy my photography. The last question now. This is something that you probably have done more than a hundred times, probably thousands of times, but this is one of those questions that young scientists and actually those that have careers that didn't make that switch to the science, but really wish they had and might want to do it later on. So the question is what advice do you have for a young scientist or perhaps someone who always wanted to go into science as a career? 



I think that there are two things that you have to deal with If you really want to get into science. You have to be tenacious, you have to be willing to make mistakes and bump your head against the wall and do it again and again, and you have to have passion. Passion is the driver more than anything else. You have to have a passion for discovery and then you have to have the tenacity to work through the failure. 

Dr. Biology:


I couldn't agree more. I also had one more thing, I say easily amused, in other words, the smallest things that you notice and observe, you can revel in, because sometimes it's those small things that make the big difference. 



I study societies of 40,000 individuals that occupy a fairly large space. There are people who study teeny, teeny, tiny things, where they look at them under the microscope, and they're just as excited about what they do as I am about what I do. 

Dr. Biology:


Well, Rob, I want to thank you again for being on Ask a Biologist. 



Well, thank you for having me. 

Dr. Biology:


You have been listening to Ask A Biologist, and my guest has been Robert Page. His research with Honeybees has spanned over four decades. To learn more about these amazing insects, be sure to look at the episode notes and chapter links. And for those who would like to dig deeper into Honeybees, Rob also has an online course based on his book Art of the Bee. You can take that course from just about anywhere, including home, or if you don't have time for a formal course. The videos are also on YouTube


The Ask A Biologist podcast is produced on the campus of Arizona State University and is recorded in the Grassroots Studio housed in the School of Life Sciences, which is an academic unit of The College of Liberal Arts and Sciences. And remember, even though our program is not broadcast live, you can still send us your questions about biology using our companion website. The address is, or you can just use your favorite search tool and enter the words Ask A Biologist. As always, I'm Dr Biology and I hope you're staying safe and healthy.

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Bibliographic details:

  • Article: Secrets of the Honeybee
  • Episode number: 131
  • Author(s): Dr. Biology
  • Publisher: Arizona State University School of Life Sciences Ask A Biologist
  • Date published: December 11, 2023
  • Date accessed: June 12, 2024
  • Link:

APA Style

Dr. Biology. (2023, December 11). Secrets of the Honeybee (131) [Audio podcast Episode.] In Ask A Biologist Podcast. Arizona State University School of Life Sciences Ask A Biologist.

American Psychological Association. For more info, see

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Dr. Biology. "Secrets of the Honeybee." Produced by Arizona State University School of Life Sciences Ask A Biologist. Ask A Biologist Podcast. December 11, 2023. Podcast, MP3 audio.

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"Secrets of the Honeybee." Ask A Biologist Podcast from Arizona State University School of Life Sciences Ask A Biologist, 11 December, 2023,

Modern Language Association, 7th Ed. For more info, see
The Art of the Bee YouTube channel

If you like this podcast episode and would like to learn more please visit The Art of the Bee YouTube channel. There you can explore the world of honeybees with Robert Page as your guide.

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