Bee Domestication

This article is one big paraphrase of a paper I wrote for a class I took with Mark Feinstein at Hampshire College called Cognition in Domesticated Animals in 2012. I highly recommend to course to anyone if it is still offered. For the sake of the audience I have cut down the length of the original paper. However I have also added extra information on domestication in order to contextualize some of the information provided. If there are any glaring problems with this article, please contact me. I have also maintained all of the citations in case you are interested in reading more about this topic. Where information was added I included a link for further information or the full citation in the text.

Bees communicate the location of flowers and food by "dancing". Wikipedia now has a great, well cited article on this behavior. This was first noted by the Austrian scientist Karl von Frisch in his book Die Tanzsprache der Bienen (English: The Dance Language and Orientation of Bees). Today our understanding of how bees communicate through "dancing" is much more better understood and documented. The direction of the bees' "dance" is in direct relation to the hive and the position of the sun in the sky. It is mostly believed that bees interpret another bee's dancing as s worker honey bee’s brain contains about 850,000 cells, half of which belong to the occipital lobe (half their brain is used for vision.) (Menzel 191 - 201). Vision is incredibly important for bees in general, and like birds, they can see ultraviolet light.  Recent research shows that bees cannot communicate this way in the dark. We also now know that the duration of the part of the dance where the bee moves a certain way (zigzags) signifies distance of the location in question.

15,000 year old cave painting
found in Valencia, Spain.

Humans and bees have always coexisted as far as we know. Before beekeeping, hunter-gathers would climb trees in order to retrieve honey from nests. Wall paintings showing this behavior have been found in countries on different continents across the globe including Zimbabwe, China, and Spain. The presence of these artifacts across the world suggests that the means by which people collected honey from bees developed convergently around the same time in different societies. One wall painting in Spain shows how humans would put giant ladders against trees in order to climb up and collect the honey. In Thailand permanent ladders made out of bamboo would be installed into the trees in order to access the honey bees’ nests (Oldroyd 213). In Vietnam, Apis Dorsata honeybees were kept in rafters. African tribes would, and still do, create artificial “hives” out of suspended wood logs, which could be placed near a dwelling. Hieroglyphics show us that the Egyptians kept hives in clay pots, as did the Greeks.  Later, in the seventeenth century, the Greeks started using wicker hives instead of clay. Meanwhile the Romans had a type of hive made of wicker, oak, and dung. When the Romans invaded Briton around 45 AD, they brought the practice of beekeeping with them.

Aristotle was one of the first people to write about the inner workings of beehives. Pliny the Elder documented that The Romans were likely the first to develop transparent hives from horn of lantern and "mirror stone" to study bees. Virgil too wrote concerning bees and where to place apiaries. He was the first to document that there was a class structure within a bees nest. Some oversights made was that he assumed the queen bee was a king. He also remarked that pollen stuck to bees in order to act as a ballast for when they flew. Some countries today such as Oman and Yemen still refer to the Queen Bee as a sheik, meaning they consider it a male (Free 100). 

When we start thinking about domestication, we need to think about what that word really means. Is it something that we can measure? Today we know that some animals can be tamed and others cannot. That there are some animals that could have been tamed, but are very much wild. Plenty of people have barn cats for example. But what does the process of domestication look like over history and how did it develop? In this case it's best to talk about dogs since they're the best example when it comes to domestication.

One of the theories that have been suggested as an explanation for the domestication of dogs is self domestication. Dogs that had a smaller flight distance (the measurable distance a person could get to a dog before it runs away) would eat food scraps from neolithic "garbage dumps" humans were making. This reduce in flight distance may be the result of an altered state of brain chemistry, which in turn would affect other behaviors. For example: while humans would eventually select and breed for specific behavioral and physical traits in dogs, there would be other measurable traits that would not initially be accounted for; the size of a domesticated dog's skull and brain is on average smaller than their wild type (Serpell, James (1995). The Domestic Dog; its evolution, behaviour and interactions with people. Cambridge: Cambridge Univ. Press. p. 35. ). It is important to delineate that a bigger brain does not make a smarter brain.

Further examples of differences between domesticated animals and their wild types::

This strange link between coat color and temperament stems from a relationship between pigment production, hormones, and neurochemistry. It is not the case that coat color causes a difference in temperament, but rather that certain physiological processes underlie facets of both coat color and behavior. In particular, the hormones and neurotransmitters involved in the stress response and other behaviors are closely integrated with pigment production.
For example, the neurotransmitter dopamine and the hormones noradrenaline and adrenaline, which are involved in the stress response, have the same biochemical precursor as the melanin pigments (Anonymous 1971, Ferry and Zimmerman 1964). In addition, dopamine directly influences pigment production by binding to the pigment-producing cells (Burchill et al. 1986). Dopamine indirectly influences pigment production by inhibiting pituitary melanotropin, also known as melanocyte stimulating hormone (MSH), which is responsible for stimulating pigment cells to produce pigment (Tilders and Smelik 1978).
Therefore, by breeding only the most docile animals in a group, humans select for physiological changes in the animal's hormonal and neurochemical systems, changes that impact morphology and physiology -- including fur color. A change in fur color during domestication may therefore be an incidental byproduct of selection for tameness.

It is therefore suggested that more tame domesticated animals have less melanin:

The character Mongo riding a tame, mostly white bull in the film Blazing Saddles.

 We also need to think about how we view the bee. A single honeybee cannot exist without the hive. The word eusocial has been used to describe their complex hierarchical behavior, and the beehive has been described as operating as a “super organism" - that is that the hive itself is seen a single organism itself. Due to the beehive existing as a single organism, and early man’s misconception as to how bees mated, it was tough for one to directly or artificially select for the right honey bees due to the fact that so many of their behavioral traits are linked to social interaction (Menzel 27). There are many different definitions of what it mean to be conscious. My personal definition is that to be conscious is to understand that you are a separate entity to the world around you. This means that I consider dogs that don't freak out at their reflection in a mirror as possibly being conscious. It means that I don't find individual bees as having consciousness, but I do see bee hives as being self-conscious. Alternatively, individual humans are conscious, but a physical group of people is very much not conscious. I also use the terms consciousness and self-awareness interchangeably. If you'd like to know more about super organisms, the book The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies is an excellent resource.

There are similar measurable traits found in honey bees versus other bees that are not used for honey production. The honey bee was are familiar with in North America is specifically the Italian honey bee, Apis Mellifer which European settlers brought with them in 1859. It has been suggested by sources that the Italian honey bee was selected out of all the honey bees for a verity of reasons: they occupy less space in the nest, they swarm less than even other honey bee species, they are more adaptable to other climates, and most importantly; they produce more honey than other honey bee species (Frank, et el.; 2000). Compared to their wild-types, honeybees initially produced a honey that was edible. Some wasps, such as the polistinae wasp (paper wasp) produce honey that can be poisonous to human. Over time bee were bread to select for quality of honey; texture, color, and a final product that is free of brood are all important factors (Free, 124).

The missing traits are that there are no current data or research suggesting that honey bees have smaller brains than their “wild” counterparts, or are whiter in color. This brings up a lot of interesting questions pertaining to our relationships with different classes of animals, and how that physically affects them. Bees absolutely do have hormones, so what is different about insects or bees where we wouldn't see the same changes in them over time? 

Today beekeepers are able to even more granularly select for desired traits by removing larva from their cells in the hive and placing them in “queen cups” where they can then be introduced into another colony. There are many more aspects today that beekeepers know to control for. “Cleaning behavior” which is the removing of damaged larva, and debris from the nest. Grooming is also important; this is the removing of mites from the bees themselves. A foreign mite introduced from another bee species has the potential to wipe out a whole race of bees. Bees stealing honey from other hives is another behavior that one beekeeper reports, and supposedly can be controlled for (Cushman). It's completely possible some beekeepers in the past know to select for these traits, but no such record of doing so has been found. Nest size is another factor that may have been selected for in bees. Considering more bees mean more honey, solitary bees are never used for honey cultivation. A healthy honey bee nest can have anywhere from 50,000 to 60,000 bees (World Book Encyclopedia; 154 - 161). Compare to a paper wasp nest, which can house around 5,000 wasps (Strassman). Due to the fact the nests are smaller and wasps are carnivores, they do not produce as much honey as a honey bee. Wasps also construct their nests out of paper instead of wax, which is less useful for human consumption.

[There was a section here on what might be causing colony collapse. Pesticides still continue to be believed to be the leading cause. New research also shows that moths at night play an integral role in pollenization.]

A bee on a yellow Rudbeckia hirta flower.

The pain score for honey bees on several different scales tends to fall in the middle. On the Starr Sting Scale of Pain, insects in the Apidae genus all score a two on a scale of one to four (Starr; 1985). On the Schmidt Sting Pain Index honey bees once again score a two out of four; alongside yellow jackets and wasps. The paper wasp scores a four on the Schmidt scale. However honey bees are less aggressive than wasps. Honeybees also die after stinging you, while other bees do not. For those who have been fortunate enough to never experience it, wasps bite and sting at the same time. This could have to do with the fact that honey bees consume nectar, while wasps are carnivores and therefore have to hunt and fight to survive. Online sources suggest that the bee stinger has evolved through fighting between bees, however all of these sites cite an outsourced Wikipedia article.

The Africanized honey bee for example may have adapted its aggressive behavior from having to protect their nests from predators in their environment; possums, bears, badgers, jackals and other predators that would destroy their nest. Sure enough, one of the biggest threats to bee keepers is bears destroying their nests. At UMASS the only bee hive we had that didn't get destroyed by a bear was on the roof of a building. The climate of Africa also means that the Africanized bees are more tailored to harsher climate with less water. Due to this reason, Africanized bees have been able to easily travel outside of their habitat, much to the discontent of the people who come into contact with them. Beekeepers in Brazil have successfully been able to breed Africanized bees into their apiaries and over time, select for less aggression (Tew). The only major difference between Africanized bee and honey bees is that the Africanized bees are more aggressive, and will pursue a perceived threat for a longer distance (Tew).

Right now there may not be much physical evidence of the bees being domesticated, but I think that more research needs to be done looking at hives rather than individual bees themselves. Nevertheless it seems that, compared to their wild types, the “domestic” bees are much more docile. Comparison of the brain size and density of honey bees to wasps and their Africanized counterparts would prove to be interesting. 

 
Sources:
"Bee." World Book Encyclopedia. Vol. 2. Chicago: Feild Enterprises Educational Corporation, 1973. 154-61. Print.

Cushman, Dave. "Honey Bee Colony Assessment Criteria." Beekeeping & Bee Breeding. 10 May 2005. Web. 9 May 2011. <http://www.dave-cushman.net/bee/assessmentcriteria.html>.

Frank, P., L. Garnery, G. Celebrano, M. Solignac, and J.-M. Cornuet. "Hybrid Origins of Honeybees from Italy (Apis Mellifera Ligustica) and Sicily (A. M. Sicula)." Molecular Ecology 9.7 (2000): 907-21. Print.

Free, John B. Bees and Mankind. London: Allen and Unwin, 1982. Print.

Hunt, James H., Anthony M. Rossi, Nels J. Holmberg, Samuel R. Smith, and William R. Sherman. "Nutrients in Social Wasp Honey." Physiology, Biochemistry and Toxicology 91.4 (1998). < http://www.umsl.edu/~huntj/Number%2064.pdf >.

Menzel, Randolf, and Alison Mercer. Neurobiology and Behaviour of Honeybees. Berlin. Springer, 1987. Print.

Oldroyd, Benjamin P. (2007). "What's Killing American Honey Bees?". PLoS Biology 5 (6): <http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1892840.>

Oldroyd, Benjamin P., Siriwat Wongsiri, and Thomas D. Seeley. "Asian Honey Bees: Biology, Conservation, and Human Interactions." Harvard University Press. Print.

Ramirez, M., E. Rivera, and C. Ereuc. "Fifteen Cases of Atropine Poisoning after Honey Ingestion." Vetinary and Human Toxicology 41.1 (1999): 19-20. Print.

Starr, Christopher K. "Pain Scale for Field Comparison of Hymenopteran Stings." Journal of Entomology 20.2 (1985): 225-32. Web. 10 May 2011. <http://www.ckstarr.net/cks/1985-PAIN.pdf>.

Strassmann, Joan E. "Social Behavior of Polistine Wasps." Rice University Web. Rice University, 1 Nov. 2006. Web. 9 May 2011. <http://www.ruf.rice.edu/~evolve/Waspweb/wasphome.html>.

Tew, James E. "Africanized Honey." Ohioline. Ohio State University. Web. 9 May 2011. <http://ohioline.osu.edu/hyg-fact/2000/2124.html>.