The Lives of Bees

by Thomas D Seeley

  hygienic. Many commercial queen producers in the United States score

  their breeding stock using this freeze- killed brood assay, because there is

  evidence that hygienic behavior is an important mechanism of resistance

  to Varroa destructor, just as it is to American foulbrood and chalkbrood.

  Colonies with hygienic workers do remove more Varroa- infested pupae

  than do those without hygienic workers.

  Another successful program of artificial selection with Apis mellifera was

  conducted in the 1960s by William P. Nye and Otto Mackensen, working

  for the U.S. Department of Agriculture. These researchers produced in-

  bred lines of honey bees that ranked high and low as collectors of pollen

  from alfalfa, a plant that honey bees visit mainly for nectar and for which

  they are usually rather ineffective pollinators. In a test performed in a loca-

  tion in northern Utah with diverse sources of pollen, using bees in the fifth

  generation of selection, they found that the percentages of pollen collec-

  tors returning with loads of alfalfa pollen were 54 percent and 2 percent,

  for the high and low lines, respectively. Several commercial seed compa-

  nies in the United States extended this research. In these studies, multiple

  lines were selected and then crossed (to avoid inbreeding), followed by

  further selection and more crossing. After three years of breeding, colonies

  in the selected lines collected 68 percent of their pollen from alfalfa when

  placed in test areas containing fields planted with alfalfa, safflower, cotton,

  melons, and sugar beets, while the control colonies in the same location

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  (the check stock) gathered only 18 percent from the alfalfa. The alfalfa bee

  story is another convincing example of what can be done in the breeding

  of honey bees. In the end, though, these selected lines of honey bees were

  not widely adopted in the commercial production of alfalfa seed, probably

  because they were much less effective than certain species of solitary bees

  that are regular, and efficient, visitors of small legumes, including alfalfa.

  NO DISTINCT BREEDS OF HONEY BEES

  Despite the decisive successes in breeding honey bees for hygienic behav-

  ior to strengthen their resistance to American foulbrood, chalkbrood, and

  tracheal and Varroa mites, and in breeding honey bees for alfalfa pollination

  behavior, there is no evidence that artificial selection has altered in any

  general way the behavior of honey bees. Why is it that the breeding of

  honey bees has had few, if any, strong and lasting effects? Certainly, it is not

  for lack of variation among colonies in traits relevant to beekeeping. Bee-

  keepers know that some colonies glue everything together in their hives

  with propolis, while others hardly use it all; that in some colonies the bees

  run about when the hive is opened, while in others they stay calm (Fig.

  4.5); and that some colonies sting fiercely when disturbed, while others

  are much less defensive. Moreover, when we look across the honey bees

  living in their homelands in Europe, we see variations in color, morphol-

  ogy, and behavior associated with geography—the differences among Ital-

  ian bees, Carniolan bees, Caucasian bees, Irish black bees, and so forth—

  but we do not see distinct breeds of honey bees.

  How different this is from the other animals that are hugely important

  to humans. Consider dogs. Like modern honey bees, modern dogs are the

  descendants of ancient European animals, specifically gray wolves ( Canis

  lupus), but unlike modern honey bees, modern dogs ( Canis familiaris) have

  been profoundly shaped by their domestication. This is clear from the di-

  versity in form and behavior among their breeds, for example: German

  shepherds, beagles, dachshunds, Labrador retrievers, Chihuahuas, Irish

  wolfhounds, Scottish terriers, and dozens more. The same is true for

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  Fig. 4.5. Worker honey bees (and several drones) standing calmly, and still dis-

  tributed evenly, on a comb that has been pulled from a hive.

  modern cattle, which are the domesticated descendants of the stately,

  reddish- brown, long- horned wild oxen (aurochsen, Bos primigenius) that

  we know from Ice Age cave paintings. From this ancient species, which is

  now extinct but was still alive in Roman times and was described by Julius

  Caesar as “little smaller than an elephant,” we have bred modern cattle to

  give us meat, hides, and milk, as well as muscle strength and stamina for

  pulling plows and wagons. And as with dogs, the effectiveness of artificial

  selection in cattle cannot be doubted, given the diversity in form and func-

  tion among the breeds of modern cattle ( Bos taurus): Holstein- Friesian,

  Belted Galloway, Texas Longhorn, Brown Swiss, Black Angus, Scottish

  Highland, and hundreds more.

  Why is it that bee breeders, unlike dog breeders and cattle breeders,

  have changed Apis mellifera so little over the past 10,000 or so years? Part

  of the answer is that we have had the full tool kit for breeding honey

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  bees—artificial queen rearing and instrumental insemination—for less

  than 100 years. What is probably a much bigger part of the answer, how-

  ever, is that the tools now available for artificial selection with honey bees

  have not been used widely and persistently. Indeed, I suspect that in most

  places where Apis mellifera lives in Europe and North America, the effects

  of artificial selection are minimal because most queen bees mate outside

  of human control: high in the air and with whatever drones they encoun-

  ter. Many, perhaps most, of these drones will come from colonies living

  in trees, buildings, and the hives of beekeepers who do not manipulate the

  genetics of their colonies. This situation means that any changes in the

  genetics of honey bees created by the work of bee breeders will, over

  time, be erased. It also means that, in many (perhaps most) places, the

  genetics of honey bees is shaped far more by natural selection for traits

  that boost the genetic success of colonies living on their own than by ar-

  tificial selection for traits that can boost the profits from colonies owned

  by beekeepers. This explains why it is that honey bees do not need our

  manufactured hives, and instead are still perfectly at home in a hollow

  tree. Indeed, most beekeepers have watched with dismay how readily their

  bees will abandon the former for the latter during the swarming season.

  The step back to living in the wild is but a short one for the bees housed

  in our hives.

  APIS MELLIFERA, A SEMIDOMESTICATED SPECIES

  While we humans have successfully manipulated the genes of maize, dogs,

  and cattle, we have not made fundamental changes in the genes of honey

  bees. We have, however, made great use of our second basic way of boost-

  ing the gains from our plant and animal partners: manipulating their envi-

  ronments. The first stage of doing so with
honey bees occurred thousands

  of years ago when we induced colonies to make their homes in our hives.

  This gave us control over where honey bee colonies live, and it enabled us

  to reach into their homes and take from them what we wanted: honey-

  combs and beeswax. These days, the technology of apiculture has advanced

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  to where beekeepers have many sophisticated means for controlling the

  living conditions of their colonies to raise their production of the goods

  (honey, pollen, beeswax, royal jelly, and occasionally venom) and the ser-

  vices (pollination) that we seek from honey bees.

  The most basic of all the environmental manipulations performed by

  beekeepers to boost the profits from their colonies is to move them to

  where they will make a valuable honey crop or where a fruit or vegetable

  grower needs their pollination services. Many beekeepers in Scotland, for

  example, move colonies to heather moorlands in August to get crops of

  the richly aromatic, reddish- orange honey made from the nectar of the

  ling heather ( Calluna vulgaris). This is the Rolls- Royce of honeys, so it is

  profitable to transport hives up to the purple- heather- covered hills in au-

  tumn (Fig. 4.6). Similarly, some beekeepers in New York State will move

  colonies to large fields planted with buckwheat in hopes of getting crops

  of the dark and peculiar- flavored but prized honey this plant yields. I know

  one beginner beekeeper who, unfamiliar with the odor of the buckwheat

  nectar that his bees were collecting, feared that something had died near

  his hive.

  The most impressive, indeed mind- boggling, example of beekeepers

  moving their colonies for pollination service is the trucking of some 1.5

  million colonies—more than half of all the colonies in the United States—

  to the almond groves in the Central Valley of California. This involves in-

  tense management of the bees. Even before they are loaded onto an ar-

  mada of tractor trailers for their cross- country trips to California from

  places as distant as Florida and Maine, many colonies are fed sugar syrup

  and pollen patties to stimulate their brood production so they will meet

  the colony- size requirement of their pollination contracts. Once the colo-

  nies reach California, in late January or early February, they are dispersed

  among the 325,000 hectares (800,000 acres) of almond orchards that

  stretch from Sacramento to Bakersfield. When the almond bloom is over,

  in early March, some beekeepers will truck their colonies north to the

  apple and cherry orchards in Washington State to fulfill more pollination

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  Fig. 4.6. Two hives of honey bees that have been taken to a heather moorland in

  the Scottish Highlands. The purple hillside in the background, near Nairn, is

  covered with ling heather ( Calluna vulgaris) in bloom.

  contracts, while others will head east to make crops of honey from the vast

  fields of alfalfa, sunflowers, and clover in North and South Dakota.

  A beekeeper can increase the income earned with his or her colonies

  by manipulating not just where they live but also how they live. For instance,

  in a natural nest, a colony’s beeswax combs are firmly attached to the

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  Fig. 4.7. Wooden frames holding combs filled with honey in a honey super that

  is sitting atop a hive.

  ceiling and walls of the cavity, but in a modern hive the bees are induced

  to build their combs in rectangular frames made of wood or plastic that

  hang like folders in a filing cabinet inside the stack of rectangular wooden

  boxes that form a hive. This arrangement makes it easy for the beekeeper

  to remove honey- filled combs by pulling the frames holding honeycombs

  straight up and out the top of a hive, rather as a file clerk extracts a folder

  (Fig. 4.7). It also makes it easy for the beekeeper to inspect (or leave un-

  disturbed) the brood- filled combs—those in which the queen has laid

  eggs and young bees are developing—which rest in frames hanging in the

  lower boxes of a hive. Another way that a beekeeper strongly manipulates

  the activities of his or her bees is by inserting into each frame a founda-

  tion, a sheet of beeswax (or plastic) that has the hexagonal pattern of

  honey bee comb embossed on each side. When the bees come upon a

  sheet of foundation, they take it to be the start of a comb and they com-

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  plete it, or as beekeepers say, they “draw it out.” In doing so, the bees build

  the new comb following the hexagonal cell pattern on the foundation.

  This usually guides the bees to build combs whose cells are of the smaller

  size used for rearing worker bees, not the larger size needed for rearing

  drones. In this way the bees are inhibited from rearing drones, and this

  boosts their colony’s honey production. It also, however, reduces its re-

  productive success.

  Besides the invention of tools like movable frames and comb founda-

  tion, the craft of modern beekeeping rests on sophisticated methods of

  colony management. Perhaps the most important one is the control of

  swarming. Beekeepers achieve this primarily by housing their colonies in

  large hives so the colonies do not become overcrowded. Some beekeepers

  will also insert frames with empty comb between the frames with brood-

  filled comb, to prevent congestion in the central brood- nest part of each

  colony’s nest. A few will even cut out queen cells—the special cells in

  which new queens are being reared—to disrupt this essential step in a

  colony’s preparations to swarm. By preventing swarming, the beekeeper

  forces the colony to invest less in its reproduction (fewer swarms) and

  more in its growth and survival (more combs, more bees, and more honey)

  than it would if it were left alone. Table 4.1 lists the principal tools and

  techniques used in modern beekeeping to boost colony productivity.

  Because beekeepers have not made fundamental changes in the genetics

  of their livestock, in the way farmers and herders have done with their

  cattle, sheep, chickens, and other farm animals, it is a mistake to include

  the honey bee in the list of animals that are truly domesticated. At the same

  time, however, because beekeepers strongly manage the environments in

  which their honey bee colonies live, both on the landscape scale (by truck-

  ing colonies about) and on the beehive scale (by managing the bees’ living

  quarters), it is a fact that the honey bee is not a totally wild species. I sug-

  gest, therefore, that we consider Apis mellifera a semidomesticated species,

  one whose genetics we have changed very little but whose environment

  we can, and often do, change very much. I also suggest that we recognize

  that even though we have pressed millions of colonies of this industrious

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; Table 4.1. Key tools and techniques used by modern

  beekeepers to boost colony productivity

  Tool

  Effect

  Movable- frame hive

  Easy manipulation of colony and its nest’s contents

  Comb foundation

  Strong control of location and type of comb in hive

  Queen excluder

  Clear separation of brood from honey stores in hive

  Section boxes for comb honey

  Honey combs are built in ready- for- sale containers

  Queen cages

  Easy transport and shipping of queen bees

  Centrifugal honey extractor

  Efficient spinning of honey from combs

  Honey uncapping knives and machines

  Efficient uncapping of honey combs

  Smoker

  Powerful calming of bees

  Artificial queen cells

  Mass production of queen bees

  Bee escapes/boards

  Easy removal of bees from honey combs

  Chemical repellents/fume boards

  Easy removal of bees from honey combs

  Feeder frames and pails

  Easy feeding of bees to stimulate brood production

  Pollen traps

  Easy collection of pollen for stimulative feeding

  Pollen substitutes

  Replacement of pollen for stimulative feeding

  Medications

  Reduced levels of disease

  Bee suits and gloves

  Colonies can be worked rapidly

  Technique

  Swarm control

  Strong colonies for pollinating and honey making

  Housing colonies in large hives

  Boost colony’s investment in honey production

  Commercial queen rearing

  Mass production of queen for establishing colonies

  Stimulative feeding

  Accelerated brood production and colony growth

  Moving colonies to work sites

  More pollination work and better honey production

  bee to live under our care, near our homes, and into our service, there

  remain countless colonies of this marvelous bee living without our care,

  far from our homes, and apart from our aims. These wild colonies show

  us that honey bees have not yielded their nature to us, for whenever they

  live on their own, they still follow a way of life set millions of years ago.

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