Organic agriculture then and now

 Once upon a time I was asked to attend a meeting about business opportunities in organic agriculture. The meeting was organized by a large international chemical company that paid my bills. I was invited because I ran an innovation lab for the company to find safe pest-control alternatives to synthetic pesticides, and this was a brainstorming meeting to come up with ideas. I was certainly in the minority. My employer hired me and my lab scientists to make the company look good to shareholders. But almost all of the company’s products and employees focused on big-market chemicals. 

The meeting started with a table topped with fruits and vegetables. One of the company’s product managers (who managed a billion dollar insecticide business) challenged the meeting attendees to identify which of the foods were grown organically and which were grown using traditional industry products. After tasting the various items, the results were inconclusive. It wasn’t obvious which were organic and which were not - at least based on taste alone. A few of the participants identified some of the  organically grown vegetables because they were limp and some of the fruits were covered with a patina of mold. These items, it turned out, were intentionally added to the experiment to prove a point: you can sometimes identify organic products by how bad they look. 

I wasn’t impressed, and today (a couple of decades later) I don’t remember anything else about the meeting. What I do remember is going back to the lab and wondering if my efforts were making a difference at all, or if I, and the scientists who worked for me, were spending time on meaningless projects. What kept me from quitting was an abiding belief that reducing pesticide use, especially on the foods that we eat, was a worthwhile challenge. 

In the end, however, it didn’t make any difference. I didn’t have to quit. I was fired because I refused to reprimand one of my scientists for finding a problem with one of the company’s new insecticides. This was stressful at the time, both for me and for my wife. In hindsight, though, it turned out to be a huge relief.

It wasn’t obvious, all those many years ago, if there really was a health benefit from eating organic products. Avoiding pesticides was a good thing, for sure, but couldn’t you just wash your food and limit any residues that way? I spent many hours reading and wondering about these sorts of things. In my mind the word “organic” basically meant a food item that had not been sprayed with a pesticide and hadn't been grown with a synthetic fertilizer. And they were marginalized or ridiculed by most Americans.

Then honey bees started dying by the millions and I watched as the chemical industry started spending some real money on environmental issues (sort of). Honey bees, as we all know, are really important pollinators to many of the fruits and vegetables we enjoy, including organic fruits and vegetables. These are the crops that also receive uncounted applications of pesticides every year. And when people started claiming that the insecticides were the reason for honey bee deaths, the industry became defensive. I know because I had a front row seat to the self-serving counter-arguments for two decades. I guess it would be more accurate to say that I was on stage, playing a minor (and inconsequential) role as the court jester. 

These findings changed the way we use insecticides, especially the group of insecticides called neonicotinoids that move into flowers and can become toxic to pollinators that drink nectar. Growers try to time these applications in order to avoid spraying when bees are active or when nectar is flowing. But not all growers are equally conscientious and problems occur all the time. 

This focus on eliminating insecticides is certainly one of the key components of the word “organic” and the bee deaths that were covered frequently in the news at the time breathed new air into the organic movement, for sure. But I was beginning to realize that the importance of the word was much broader than I originally imagined. 

There are a handful of definitions for the word “organic.” In my dictionary the third definition is the one we think of in an agricultural context - referring to food grown without pesticides and artificial fertilizers. The first definition refers to any organ system of the body. My lungs, following this definition, are organic. But it is the second definition that should be our primary focus when discussing organic agriculture. My dictionary defines it as follows: “Of, pertaining to, or derived from living organisms.” And the reason I think this is the better definition is because there are uncounted millions of living organisms that go into making organic foods so healthy and densely packed with nutrients. In fact these uncounted microorganisms are now known to be the major difference between organic and non-organic foods. If I were to go back to that meeting decades ago. I would tell the group that a field trip was needed. We would need to look at the soil in which the different food items were grown and see which soils were more alive. And then we would start to understand how it is that these microorganisms are able to break down all kinds of molecules in the soil to a form that plants can take into their roots - and ultimately make available to us.

The reason we need to limit pesticides is so that living things can flourish. I don’t mean that the insect pests that eat vegetable leaves need to flourish, nor that the many diseases that destroy roots should, and I’m certainly not wanting to encourage invasive weeds that choke out our seedlings. I do mean that the goal of organic agriculture should be the flourishing of living organic beings - including those that live in the soil and that create an environment for healthy plants.

Pesticides are an important challenge in our need to feed the world, but they are just one piece of the challenge. Equally daunting is the fact that our mass-produced foods lack many of the plant nutrients that we need for lasting health. Organically grown foods have a higher content of these healthy phytochemicals than do foods grown with many artificial inputs. This important fact is only now starting to sink in. 

This has not always been clear because we tend to judge food by what it looks like on the outside. And, as my industry experience taught me, applying pesticides and fertilizers does, in fact, produce bigger, more abundant, and showier crops. So while it is true that the healthy phytonutrients are much more abundant in organically grown plants, there is no way to know this when looking at a grocery bin full of vegetables. There are no labels containing nutrient facts for every box of organic apples flown in from the Rodale Institute in Pennsylvania, or of fresh shallots just harvested from Filaree Farms in Washington. One has to take these facts on faith, unless you have a body (like mine) that forces you to pay attention. Many people, when they get to be my age, learn first hand about the importance of nutrient dense foods because they make us feel so much better. When we ignore them, we pay a heavy price. 

If I could go back to that industry meeting all those years ago and reply to the cynical business manager, I would also have asked for an analysis from a biochemistry lab, or a nutrition lab, comparing the micronutrients of both the organic and non-organic foods. Such a request would probably have been scoffed at. A carrot is just a carrot, after all. Or so most of us seem to think. And to be honest, that is how most of us thought about food a couple of decades ago. 

Today I cannot emphasize enough how important micronutrients are and of the role they play inside the cells of our bodies. We need to rewrite some of our beginning biology and nutrition books that focus primarily on the role of the bigger and more abundant molecules.

I can give an example of how this plays out in classrooms all around the world based on my own experience teaching beginning biology classes. Most courses (and textbooks) begin by introducing the basic chemistry of living things. We cover atoms and the various kinds of bonds that hold them together in molecules. This is all fairly basic information. We talk about the importance of the element carbon to life because it is so important in forming the basic structures of our bodies. Oxygen, nitrogen, and hydrogen are likewise emphasized. These four elements are not only essential to life, but they are also the most abundant elements in our bodies. And being deprived of these elements can lead to illness and even death in just a short amount of time. So it is appropriate to focus on them.

We also recognize that there are other essential elements that are very important as well, but less urgently so, and that we need in smaller amounts. These include: calcium, potassium, phosphorus, sodium, silicon, chlorine and magnesium. 

Then there are the trace elements that are sometimes essential as well (depending on the species) but that are present in much smaller quantities. For example, we need iodine for proper hormone function and we need iron in order to capture oxygen and remove carbon dioxide from our bodies. We just don’t need very much.

In short, we do a pretty good job of teaching the importance of the basic chemistry of life, but with a hitch. It often comes across implying that each of these elements acts alone, or in ways that should be understood in predictive algorithmic ways. In other words, we ignore the many molecular interactions that occur every moment inside our cells and we generally fail to notice the many phytochemicals that only organic foods supply. We tend to overlook them because they are complex and are considered too difficult for beginning biology students. These molecules need a broader treatment in a more challenging chemistry class, or so we assume.

Most students leave college believing that they need to eat more fruits and vegetables and take vitamins. This is their basic understanding of what nutrition means. And of course, this is a good start. If you ask the same individuals about the importance of phytochemicals (plant-derived molecules) they will likely have no idea what you are talking about. And yet this is the big contribution of organic foods. In study after study, it is these plant-chemicals that show up more abundantly in organic produce that haven’t had soil microorganisms fried by pesticides and synthetic fertilizers. 

David Montgomery and Anne Bikle put it this way: The “consistent differences between conventional and organic crops fall out most clearly in their phytochemical and antioxidant content - that is, in the levels of compounds that halt the start of or reverse ongoing disease processes but are not generally considered nutrients” [italics mine].

Our students leave school very happy to know that nitrogen, potassium and phosphorus are all being dutifully added to our foods via synthetic fertilizers. It never crosses their minds that this is the same old story we have been telling ourselves for over a century - or maybe it would be better to say that it is the same old partial story we have been telling ourselves. 

I am holding in my hands a copy of the 1939 Yearbook of Agriculture titled Food and Life. Like most of these annual tomes from generations past, it is timeworn and largely forgotten. Like old Readers Digests condensed books, these yearbooks sell for a dollar or two in thrift shops and largely collect dust. Yet I find them interesting. They are a window into the farm life of my grandparents, and they show our agricultural priorities from a simpler time. They also bring a realistic perspective to the long history of America’s adoption of industrial agriculture. 

There are long chapters here about basic nutrient requirements. But what surprises me is how similar many of the recommendations are to our modern nutrition textbooks. All the major and minor elements are here. And while there are insights that are missing (because they haven’t been discovered yet) I could assign one or more of the chapters to my freshman students and not go far afield. 

Several of the authors point out the fallacy of perceiving healthy diets as a simple summation of calories. Apparently this was the bandwagon of the new science of nutrition during the 19th Century right after we discovered that we could measure energy stored in foods by how much heat they could produce - or literally how much of a particular food, when burned, would raise the temperature of water 1 degree Celsius. This was an important finding, for sure, and it helped us understand the relative importance of some foods in fighting starvation. 

The authors point out, however, that proper nutrition is much more than just counting calories. You have to pay attention to the basic elements that confer health. You can get all the calories you need and still get sick because you lack certain basic vitamins and minerals. Reading this is a bit frustrating because there are still many among us today that are still stuck in the counting calories mindset. 

It is also frustrating because we are making the same summation mistakes today. The difference is that we are now stuck in the count-your-vitamins mindset. We know that trace elements and minerals are important, and we tell our kids to eat their vitamins, but we let big business push these nutrients on us as if it is quantity that matters. It is this summation mindset that makes it impossible for many to understand the importance of organic foods. It also hides the importance of individual molecular contributions. 

Take hemoglobin, for example. It is a critically important molecule that circulates in our blood and carries oxygen. It is primarily made of the common elements of carbon, hydrogen, oxygen, and nitrogen. There is also a single atom of iron tucked away in the folds of these other elements and it is this one atom that gives hemoglobin its unique shape and makes it possible for it to do its job. So while it is true that you can add up all the atoms and pretend to know what is required to make hemoglobin, it is really only iron that becomes critically important - or we might say that iron is the limiting element in making hemoglobin. It might also go without saying that iron is only needed in very small amounts. 

In fact a diet with too much iron will upset the stoutest stomach, and in even larger amounts it can lead to organ failure, seizures, comas, and even death. Iron is critically important and it is also toxic - just like every other vitamin and mineral that we need in order to remain healthy. And this is the piece that we miss when it comes to the secondary chemicals produced by plants that grow in the wild or that are grown in rich organic soil. They are all handled by our bodies in a healthier way when we consume them in the context of real food, not as supplements or as fortified, refined, and packaged alternatives with endless shelf-lives. 

Plant-produced molecules are often bitter. If you try feeding them to children they pucker their lips and refuse to eat. This is particularly the case for children (and even grown-up children) that judge all foods based on how sweet they are (because they have been fed sweets by their parents from a very early age). But here’s the big surprise: many of our cells have receptors for absorbing these bitter-tasting plant chemicals. What I mean is that our cells have evolved to need some of these molecules, and when we don’t get them, our cells stop working the way they are designed to work. 

At the turn of the 21st Century, about when I began my frustrating career in industry, papers started appearing in scientific journals around the world suggesting that some plant compounds regulate cellular activity in the animals that eat them (Jacobsen et al., 2002; Lee et al., 2004). Flavonoids are an example. They are molecules found in many wild and organically grown plants that are produced as deterrents to herbivores. We now know that they also slow down cellular activity, even cellular division, and possibly slow down cancer cell activity in animals that eat them. Other phytochemicals seemed to be doing the same thing. The compounds were docking at the cell surface and sending signals that “modulated” internal activity. The abbreviation scientists came up with to describe this phenomenon was SKRM (short for selective kinase response modulators).

Jeffrey Bland summarized these findings as follows. “The studies confirmed that there are substances in certain foods that send messages to our genes through the influence of the kinase regulatory network, and that these messages control the production and release of inflammatory mediators.” The message that is slowly dawning on nutritionists around the world is that plant compounds slow down our constantly active cells. They reverse the many manifestations of inflammation, and it even seems that they can slow down inappropriate cell division. 

I no longer flinch when someone criticizes my organic choices. The benefits are much more real to me now than they used to be. Just like I felt decades ago, I’m still happy to be lowering my consumption of pesticides. But more importantly, I benefit in tangible ways from the denser phytonutrients that organic foods provide. These are nutrients that don’t get written on food labels because we haven’t figured out how to do that well yet. In the meantime, I keep planning next year’s garden even though I’m writing these words in November. Being healthy, after all, is a year-long effort. 

References

Bland, Jeffrey S. (2014). The disease delusion, conquering the causes of chronic illness for a healthier, longer, and happier life. Harper Wave. 

Jacobson, K. A., Moro, S., Manthey, J. A., West, P. L., & Ji, X. D. (2002). Interactions of flavones and other phytochemicals with adenosine receptors. Flavonoids in cell function, 163-171.

Lee, L. T., Huang, Y. T., Hwang, J. J., Lee, A. Y. L., Ke, F. C., Huang, C. J., ... & Lee, M. T. (2004). Transinactivation of the epidermal growth factor receptor tyrosine kinase and focal adhesion kinase phosphorylation by dietary flavonoids: effect on invasive potential of human carcinoma cells. Biochemical pharmacology, 67(11), 2103-2114.

Montgomery, David R. and Anne Bikle. (2022). What your food ate, how to heal our land and reclaim our health. W.W. Norton & Company. 

USDA. (1939). Food and life, yearbook of agriculture 1939. United States Government Printing Office, 76th Congress, 1st Session. 

Utah and Her Honey Bees

 The state insect of Utah is the honey bee. This is, perhaps, a trivial fact, and one that only an entomologist like me might care to remember. But even though other states (15 to be precise) have chosen the same insect, the story in Utah is unique. We care about these bees as others do. We enjoy eating their honey and benefitting from their pollinating services. But here, the honey bee also looms large as a symbol of a hard-working community surviving in a desert landscape. 

Other places typically decided on their state insects by committee and legislative processes many years after they became states. In Utah, the beehive was a symbol long before statehood was granted. In fact it was a symbol long before there were any state insects at all. The evidence suggests that in Utah it wasn’t a committee that made the decision either. Brigham Young may have come up with the idea himself sometime between July 24, 1847 and July 24, 1848 - the first year that the pioneers lived in the Salt Lake Valley. 

The odd thing about this story is that Utah and her people are not all that interested in honey bees. We certainly have beekeepers and apiaries, and we gather and sell a variety of honeys. We even have a bee lab in Logan where over a million bees are kept in small white unit trays within glass-topped drawers. But these bees are mostly native bees of the Western US. Less than 1% are honey bees. If you ask the majority of Utahns if they know the difference between a honey bee and native Anthophora bee, they would only shrug their shoulders. I know because I do this sort of experiment on a regular basis (not to be cruel, but to make a point). 

And as far as the honey is concerned, we produce less than other states that have never recognized the honey bee as a state insect. My copy of the 2002 Agricultural Statistics has Utah ranked number 23 in the nation in the number of honey producing hives. A skeptical Utahn might say that the low ranking is due to the small population of the state. But the numbers don't follow. Of the five states bordering Utah, only Nevada has fewer hives than Utah. Even sparsely populated Wyoming has more than we do.   

The modern entomologist might find these conflicting points confusing. More so because the beehive symbol shows up all over our state. The proper noun Deseret is perhaps the most obvious example. It was the name of our state before Utah was officially accepted into the Union (as the State of Deseret). It is the name given to bees in the Book of Mormon. The reference comes in the Book of Ether (3:2) where a group of Old World emigrants are traveling to the Americas. The account briefly mentions the conveyance of honey bees to the New World. 

But this reference is a mere sentence in a sacred text consisting of over 500 pages. There is nothing here that would lead anyone to assume an elevated iconic status for an insect. And yet the iconic status persists. We don’t call our state Deseret anymore (at least not very often). And since becoming the 45th state of the Union (in 1896) we are no longer known as the Utah Territory. But Utah is still very much the Beehive State and our flag proudly flies with the image of an old-fashioned hive (a straw skep) front and center.  

In his master's thesis, Michael Hunter argues that the beehive symbol has a long history that begins in monarchical Europe and develops through republican revolutions, and Masonic ritual. In Utah this evolution also included theocratic renewal. It isn’t the insect that matters, he argues, it is the old symbol of an ordered society, of hard work, and of selfless service that impressed itself upon Brigham Young and the Utah Saints. What I find so interesting is that the beehive symbol itself has endured even though our understanding of honey bee biology has changed a great deal in the last 500 years. 

A few hundred years ago we thought that a colony of bees was ruled by a benevolent king bee (Crane, 1999). A king bee was like a divine ruler, wrote Thomas Aquinas: “Among the bees there is one king bee and in the whole universe there is One God, Maker and Ruler of all things.” “The king does not sting,” was an important bee motto of the Renaissance (“Rex Spicula Nescit”, literally translated, the phrase means: “the king does not know how to sting”).

Even before Aquinas there is a long history of assuming queen bees to be kings. Bees were the symbol of kingship in Lower Egypt. And in Ephesus the officials of Artemis were called Essenes (not to be confused with the Jewish sect of Essenes) - a name that is said to refer specifically to the king bee (Hunter, 2004). Some ancient cultures (including Egypt) believed that souls took the form of bees and were watched over by a benevolent male ruler that was partially divine.

By 1609, it had been established that the large bee in the center of beehive activity was actually female. But instead of losing the royal symbol, Europeans just changed the sex of the ruler. He wasn’t a king. She was a queen. Charles Butler, an English rector, wrote that “The Bee-state is an Amazonian or feminine kingdom.” Even so, comparisons continued to be made between bees and kings. 

When the Anglo-Dutch philosopher Bernard Mandeville wrote his Fable of the Bees in 1714, he was not worried about whether bees had kings or queens. Bees were a symbol of social mores writ broadly and his purpose was to show that republican individualism should be the basis of a good society - kings were a burden. The real work of society was performed by the workers, and without the worker bees, the queen would die. 

The ideally functioning hive with its dedicated working class made its way into freemasonry as a symbol of fraternity. For the Shakers it was a symbol of religious community. For the Catholics the beehive was a symbol of a formal church with its well-functioning parts.

As 19th-Century members of the Church of Jesus Christ of Latter-day Saints were pushed out of Ohio, Missouri and finally Illinois, their views of American republicanism developed along different lines than those of either the northern or the southern states. On one occasion Joseph Smith referred to the restored church as a theodemocracy.  Joseph’s successor Brigham Young believed that a theocratic community could exist within the American republic. And it was Brighma Young who established the beehive as Utah’s foremost symbol even as the rest of the country recognized the same insect in different ways.

Val Brinkerhoff argues that the beehive in Utah represented the Kingdom of God. And it was usually depicted underneath the protected talons of our national bird - the bald eagle. These combined symbols (of raptor and hymenopteran) are consistently portrayed together in wood carvings, public monuments and even in church conferences during the administration of Brigham Young.

The truly remarkable part of this history (at least for me) is that the beehive remains a very visible and public symbol in Utah even though it no longer works symbolically as it used to. Our knowledge of bee biology has made some of the earlier symbolic notions unworkable. That said, we are still concerned about bees, at least symbolically. The recent pandemic of colony collapse disorder has been an interesting contrast. The disorder certainly worried the world. Utahns, however, weren't unduly concerned for the bees. It was always the beehive as a symbol that was important and not the bees themselves. 

But the modern interest in honeybees worldwide has to do with their biology and not with their social structure (and its symbolic history). In Utah, however, the symbolism remains. We don’t think of honeybees as ruling monarchs nor as harbingers of a future theocracy. But we do continue to see them as examples of industry, of selfless service, and of an ordered society. It is convenient (and probably proper) to ignore the fact that honeybees, for all their social adaptations, also maintain a caste system. 

I only bring this up because my ancestors were an independent group of tough desert farmers and ranchers. They knew first hand the importance of community but they were also individualists and trusted their own judgment before that of anyone else. They would never have submitted to a caste system, nor was it a part of their faith. I know this from a lifetime of experience belonging to an extended family that hails from very rural and independently-minded sagebrush communities in Utah. 

The Great Plains and the American Midwest may have experienced long periods of conflict between farmers and ranchers. But out here in Utah, by the end of the 19th Century, existence depended on a hardscrabble determination to raise food in whatever way was possible. My ancestors farmed and ranched. And they contracted out their services to locals and governmental employers whenever opportunities presented themselves. 

This combination of pluck and community does have a counterpart among honeybees. Individual worker bees may seem to go about their various jobs as subservient automatons while maintaining the hive, but when they set out on their own to build a new hive, they work very democratically indeed. 

This happens when the old queen can no longer keep new queens from developing within the hive. She becomes restless and gathers a few thousand worker bees to follow her away from the old home and swarm to a nearby branch or other available outpost. Then over the next several hours or days, scout bees go in different directions looking for a suitable place to start over. The decision on the best location is made by the hive as a group. It isn’t a decision made by the old queen nor by a select committee. It truly is a democratic decision (Seeley, 2010).  

The so-called caste system of bees comes from the 20th Century definition of eusociality which states that eusocial organisms: 1) cooperate in rearing young, 2) that there exist overlapping generations within the hive, and 3) that there is a division of labor between reproductive and nonreproductive groups (sometimes called castes). A few authors (including the well-known entomologist E. O. Wilson) have argued that humans display a weak form of eusociality as well. And this might imply that the Utah fascination with honeybees stems from a deeper willingness to accept a sociality of castes. 

Such claims are historically uninformed and misunderstand the self-reliance that is a deep part of our Utah psyche. It was Suzanne Batra (in 1966) who introduced the word “eusocial” in her work on nesting behavior in some sweat bees (very distant relatives of honeybees). And a few years later, the well-known bee taxonomist Charles Michener used the term for other bees (including honeybees) generally. Overlaying this recent understanding of bee biology onto the Utah admiration of bees (stemming from a century earlier) makes no sense. 

Neither should this argument be used for our many modern neologisms like Beehive Credit Union, Beehive Brick, Beehive Pizza; or Deseret Book, Deseret Industries, etc. Utah culture past and present, sacred and secular, biologically informed and otherwise,  is perfused with this symbol. 

Armand Mauss, in his 1994 study of Mormon assimilation, has argued that the beehive symbolism is thematically more important than just traditional industry and hard work. He contrasts the beehive with the symbol of the Angel Moroni that rises with a trumpet on top of many temples. The angel represents the spiritual and revelatory other-worldliness of the restored gospel. The beehive, in contrast, is somewhat of an excuse to be pursue wealth. It is primarily “a symbol of worldly enterprise throughout the Mormon heartland.”

My own take on this contrast is somewhat skewed, yet also informed, by an entomological sensitivity. The bees themselves should not be ignored, and they are tougher than we might expect. The ones that we have kept caged in our wooden hives are struggling to survive the many introduced pathogens and toxic chemicals that modern society continues to generate. The ones that have escaped into the wild and keep themselves in tree holes and rock cavities are doing much better. These are the ones that seem to resist our imposed “assimilation.”

I think there is a lot to be gained by the right kinds of symbols. Beehives and the social insects that live in them must certainly be counted as one of these - both for Utahns and the long apian history of many other lands. But symbols that we inherit, and hardly think about, usually fail. They can only be taken so far. The industrial behemoth that is choking our world (and that is loved by too many Utahns) seems to be looking all too favorably on our many symbolic hives. Somewhere along the way we should stop and pay more attention to the insects themselves and to their preferred wild landscapes. We have exploited them both for far too long. 

References

Batra, S. W. (1968). Behavior of some social and solitary halictine bees within their nests: a comparative study (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 120-133.

Brinkerhoff, V. (2013). The Symbolism of the Beehive in Latter-day Saint Tradition. BYU Studies Quarterly, 52(2), 9.

Butler, Charles and John Owen. (2017). The feminine monarchie or the history of bees. Northern Bee Books, Hebden Bridge (prepared from the 1623 edition).

Crane, Eva (1999). The world history of beekeeping and honey hunting. Routledge, New York.

Hunter, Michael J. (2004). The Mormon hive: a study of the bee and beehive symbols in Nineteenth Century Mormon culture. Brigham Young University BYU Scholars Archive. 

Mauss, Armand L. (1994). The angel and the beehive, the Mormon struggle with assimilation. University of Illinois Press, Urbana and Chicago. 

Michener, C. D. (1969). Comparative social behavior of bees. Annual review of entomology, 14(1), 299-342.

National Agricultural Statistics Service (2002). Agricultural Statistics 2002. United States Government Printing Office, Washington. 

Seeley, Thomas D. (2010). Honeybee democracy. Princeton University Press.