Anarchic Behaviour
Laying Workers
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Thelytoky in Honey Bees

Both the original document and I have drawn on various sources for the following information although the majority comes from the Carl Hayden document. These sources are noted at the bottom of the page, if I have missed any, I apologise.

Basically, Thelytoky is... the ability to rear workers and queens utilising the eggs from laying workers, or in some cases virgin queens. This subject has in the past been considered a rarity that only occurs in the Cape bee, Apis mellifera capensis Escholtz, but it has been found in other strains.

While Thelytoky is exhibited strongly in Capensis, it does occur in all races of Apis mellifera, but in all except Capensis, it is very rare [1]. At Capensis levels it is destructive, but at levels of one in a million it is unlikely to do much harm and on the rare occasion that the gene pool is impoverished, it can maintain genes that might be otherwise lost.

It would be wrong to consider Thelytoky as an important issue, but studying the causes and effects gives us a little more insight into the feedback that helps to regulate natural selection. It exists and is stable at it's current levels, hopefully whatever keeps it that way will counteract any selection that may be done by humans that may try to increase it.

The diploid eggs laid by thelytokious bees are formed from haploid eggs that have a modified division process in the early stages of cell division...
Below is the process as described in Capensis

Production of parthenogenetic female eggs - thelytoky Verma and Ruttner (1983) showed that the secondary oocyte fuses with a polar body in the content of the unfertilized oocyte during meiosis. This automictic mechanism was suggested by Tucker (1958) and allows no combination of loci in the offspring, unless crossing over mediates the exchange of linkage groups. Moritz and Haberl (1994) could not detect crossing over in the formation of these diploid offspring, hence all offspring of a single capensis worker are genetically identical, mother and daughter therefore form a genetic clone. Ruttner(1988) claimed that only a single recessive allele, th, at one locus determines workers to perform thelytokious parthenogenesis. The thelytokious parthenogenesis has been explained as an adaptation to the harsh, wet, windy conditions where the queen is much more on a risk getting lost during her mating flights (Tribe 1983, Moritz and Kauhausen 1984, Moritz 1986). But paradoxically, the highest frequencies of matings occurs in months in which the winds are most intense (Allsopp and Hepburn 1997). Referring to the high mating frequency detected in A. m. capensis (Estoup et al. 1994, Moritz et al. 1996, Kryger 1997) the mating risk of the queens seems not to be higher than in European races.

Taken from...
Regulation of reproductive dominance hierarchiesin Apis mellifera capensis workers Dissertation zur Erlangung des akademischen Gradesdoctor rerum naturalium (Dr. rer.nat.) vorgelegt der Mathematisch-Naturwissenschaftlich-Technischen Fakultät (mathematisch-naturwissenschaftlicher Bereich) der Martin-Luther-Universität Halle-Wittenbergvon Frau Ute Simongeb. am: 11.08.1967 in Emden Gutachter1. Prof. R.F.A. Moritz2. Prof. N. Koeniger3. Prof. R.M. Crewe.

Thelytoky does occur in the European Apis mellifera strains, but with considerably lower frequency. In queenright colonies development of most worker ovaries is suppressed by the pheromone 9 - oxy - decenoic acid and possibly other substances emitted by the queen, [2] as well as substances possibly emitted by unsealed brood. [3] Workers can develop ovaries and some can lay eggs in the absence of both queen and open brood. [4]

European laying workers generally produce unfertilized haploid eggs that give rise to drones (if they develop at all). It is rare, but there are instances of both virgin queens and laying workers producing diploid eggs and those that develop, produce true female worker or queen bees. [5]

What causes Thelytoky? First a biological mechanism is needed to produce viable diploid eggs. Secondly various natural control systems need to be bye-passed or modified {see worker policing}. The biological mechanism of how the haploid egg becomes diploid is a gobbledygook sentence that I will reproduce here...

"Cape bee workers lay unfertilized diploid eggs because during ana-phase II the egg pronucleus and the central descendent of the first polar body fuse to form a diploid zygote nucleus." [6]

But the behaviour modification is more difficult to understand!

I offer various quotes and elements of the original text that I hope will aid understanding...

"Given the high frequency of thelytoky in Cape bees, the relatively rare occurrence in domestic stocks of European bees is unexpected, since populations capable of thelytoky have an advantage over those in which laying worker eggs develop exclusively into drones." (Ruttner 1977) {This may now be better understood due to the recent work on worker policing by Francis Ratnieks at Sheffield University.}

"Without thelytoky, the survival of a colony rests completely on the successful mating of a single queen which must leave the hive to mate. If this queen does not encounter drones or does not return to the hive, a replacement cannot be produced because female larvae of a suitable age for queen rearing no longer exist, and because the first queen to emerge usually destroys the other queen cells in the colony. However, if brood from laying workers could be raised into queens the colony would have a facultative survival mechanism in case the virgin queen is lost. Thelytoky should occur with greater frequency in populations exposed to conditions that reduce the chances of a queen mating." (Moritz 1984)

"Reports indicate that in managed colonies thelytoky is expressed at a very low frequency" (Mackensen 1943). The reasons given for this statement are given below.

"Beekeeping practices inadvertently select against thelytoky. For example, swarming and supersedure can be minimized through various management techniques, and thus the possibility of a colony becoming queenless due to the loss of a virgin queen can be reduced. If colonies lose their queens and do not have brood to produce replacements, the queens often are replaced with new ones by beekeepers. Hence, there is no selective pressure for thelytoky in colonies managed in this manner. Conversely, the conditions under which the LUS strain was derived may have inadvertently selected for thelytoky. Virgin queens introduced into broodless colonies during the winter may not have been accepted by the workers in some cases, while in others the queens may not have mated or were lost on mating flights. Some of the colonies that survived may have done so because they requeened themselves with brood from laying workers. The winter requeening procedure was repeated annually using queens produced from brood of colonies that survived the previous year's winter requeening. If thelytoky was originally at a low frequency in the LUS strain at the beginning of the breeding program, the frequency possibly was increased because of continued selection followed by the production of new queens from brood of the survivors."#

"Sometimes during an inspection bees were seen biting each other, or with their abdomens in the cell assuming an egg laying position. We sampled LUS bees being bitten and dissected them to determine if they had ovary development. Whether workers assuming the egg laying position always deposited an egg in the cell also was determined."

"Once all the brood emerged in queenless LUS, CP, or cd (control) colonies, worker bees were scattered over the frames giving the colony the distinctive appearance associated with the queenless state. Upon closer examination of bees from the 4-5 frame nucleus colonies and in the observation hives sometimes workers were seen grasping each other with their mandibles."

"In a LUS observation colony, workers were seen pulling nestmates out of the cells in which they had inserted their abdomens. On other occasions, in the observation hives we saw eggs being eaten by nest mates immediately after the laying worker removed her abdomen from the cell."

"A queen produced from laying worker eggs successfully mated and produced worker and drone brood. However, eight of the nine queens produced from workers' brood either did not return to the hive after a mating flight, or were critically injured during artificial insemination."

"A honey bee colony's ability to re queen itself with the eggs of laying workers requires not only that some workers can lay diploid eggs, but that the workers can foster the cooperation from nest mates needed to construct a queen cell and rear the egg into a queen. When laying workers developed in CP or cd colonies, often queen cells were constructed and sometimes eggs were deposited inside them. However, the eggs were either cannibalized by other workers or left unattended (untended) and did not hatch."

Possible flaws in this work

  1. That there was already existent in the LUS strain abnormally large numbers of genes from Apis mellifera capensis. Such genes could have been filtering into the area of the trial over many years but at such a low rate that the colonies concerned went unrecognised.
  2. There is the possibility that the quantity of the thelytoky genes (from whatever source) was enhanced by other management, out of season breeding, the use of small celled foundation, selection of queens for shorter development times. Any or all of these may have had an effect.
  3. In the case of virgin queens laying diploid eggs there could have been a fleeting, incomplete mating that passed a small number of sperm to the queen which at later dissection were not noticed or the few sperm were "used up" in fertilising the few eggs laid.
  4. I consider it possible (but unlikely) that sperm could exist in a "loose" state from accidentally crushed drones, but I can also see many obstacles to those sperm ever being in a position to fertilise an egg, from any source, even if they survive the exposure.


[1] Onions 1912, Jack 1917, Anderson 1963, Ruttner 1976
[2] Butler and Fairey 1963
[3] Kropacova and Haslbachova 1971
[4] Perepelova 1929, DeGroot and Voogd 1954, Butler 1957, Butler and Fairey 1963, Jay 1970, Kropacova and Hasibachova 1970 & 1971
[5] Mackensen 1943
[6] Verma and Ruttner 1983

Other References...

Suomalainen 1950
The Work was carried out by... Dr. Gloria DeGrandi-Hoffman, Dr. Eric H. Erickson Jr.
Of... Carl Hayden Bee Research and Biological Control Center, Tucson, Arizona, USA
And was published 29 January 1991
The LUS strain of bee was selected by:- Delores and Edward Lusby who are commercial beekeepers working in Tucson, Arizona.

I have been selective in what I have taken from the original text, this selectivity is intended to simplify rather than mislead.


 Written... Spring 2001, Revised... 20 June 2001, Upgraded... 11 June 2006,
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