My favorite beefact™ is that each drone carries millions of identical sperm (i.e. male gametes created via mitosis); each queen only takes one maiden voyage, whereupon she typically chooses about a dozen suitors.

Her spermathacea is able to keep isolated these dozen suitors' loads, and remembers both the order and the aggressiveness of each suitor (each's "rapiness"). Queen can then choose which sperm bank to use when fertilizing each egg (i.e. creating more female workers)...

Through incredible (and largely unknown) mechanisms, queen typically develops sperm/suitor preference. Remembering the rapiness of each suitor, she can adjust hive's temperment (e.g. if skunks are robbing, be more aggressive).

As she ages, she'll run out of "The Best Sperm" (remember: it's all stored for YEARS from one mating afternoon) and have to go to B-team, C-team, etc... until the workers choose to replace her (queen mandibular hormone output also greatly reduces as she ages, which gives the hive "identity").

One day the Queen's Court will lead her to a queen cup, have her lay a fertilized egg within, ultimately sealing her fate [to be murdered by the princess queen upon her birthing, unless she is still healthy/young enough to be chosen to lead the next outgoing swarm].

Source: beekeeper that has read too many books

"Our findings indicate a fundamental genetic program that determines the neuronal circuitry and behavior of worker bees"

This dogma - that bees are little machines "programmed" by genetic code - won't lead very far. A single gene may turn off our auditory system because it controls production of a protein that's necessary for hearing, and loss of hearing would make us less social, but it's a wrong conclusion that the "auditory gene" is what "programs" our social behavior.

IMO, Michael Levin's research is closer to truth. If we extrapolate his ideas to a whole bee colony, we could find that behavior of a bee hive can be changed without messing with genes, although some of the genes make bees receptive to "social cues" that makes such large scale behavior possible.

In case you want to try honeybee experiments at home ... from the paper:

https://www.science.org/doi/10.1126/sciadv.adp3953

"Honeybee handling procedures

The honeybees were collected from Apis mellifera carnica colonies (western honeybee) at the bee yard of the Heinrich-Heine University, Düsseldorf, Germany. Female eggs were collected from naturally mated queens, which were maintained in small nuc colonies with five combs (Holtermann, Germany). To collect female eggs, the queens were caged in Jenter egg-collecting cages (Jenter queen rearing kit, Karl Jenter GmbH, Frickenhausen, Germany). For the tracking, we collected newly eclosed bees (0 to 24 hours old) from a brood comb that was maintained in an incubator at 34°C. The laboratory rearing of worker bees was done as previously reported (19). We grafted the newly hatched larvae into plastic cups (#4963, Heinrich Holtermann KG, Brockel, Germany) with worker diet # 7 (53% royal jelly, 4% glucose, 8% fructose, 1% yeast extract, and 34% autoclaved water) (74). To obtain bees with fully developed worker characteristics, we experimentally determined the amount of food provided, which was 170 μl per larvae. The larvae were kept at 34°C and 90% relative humidity. The latter was generated using saturated solution of K2SO4 (75). Before defecation, the larvae were transferred onto Kimwipe papers (Delicate Task Wipers, #066664, Kimberly Clark) and were kept in petri dishes for 2 days at 70% relative humidity, which we generated using saturated NaCl2 solutions (75). After defecation, the prepupae were separated into plates with 24 wells (#92424, Peter Oehmen GmBH, Essen, Germany) in which filter papers were placed (15 mm, grade 413; VWR, International GmbH, Darmstadt, Germany). Once they started walking, they were marked and maintained in small cages together with eclosed wt workers coming from colonies (#20104; Imkereifachhandel Jasniak, Trossin, Germany) in which water and sugar paste supplemented with pollen (#7032; Heinrich Holtermann KG, Brockel, Germany) was provided. To obtain myrGFP-mutated worker or queen bees, we reared the queens as described (17, 76). To obtain myrGFP worker bees, 12- to 19-day-old myrGFP queens were inseminated with wt drones using standard insemination techniques. The queens were treated with CO2 1 day before insemination. Inseminated queens were maintained in small nucs (“Kieler Begattungskasten,” Holtermann, Germany) with wt worker bees. The nucs were kept in a containment so that mutated animals were not able to escape into nature. To obtain newly eclosed dsxmyrGFP/+ worker bees, combs with capped cells were maintained in an incubator at 34°C."

An interesting insight led to their research (from the paper): https://www.science.org/doi/10.1126/sciadv.adp3953

(First, some interesting background:)

"The evolutionary transition from solitary to social living in vertebrates and invertebrates led to sophisticated social behaviors. During the past 50 to 150 million years of evolution, sociality in some species became so elaborate that individuals in the group forego reproduction and changed their behavioral performance to embrace collective behavior, while others specialize in reproduction leading to the development of two castes, queen and workers (1, 2). From behavioral activities of hundreds and sometimes ten-thousands of worker individuals, new properties have emerged at the collective level such as shared brood care, warfare, collective thermoregulation, nest building, and farming, which contributed to the spectacular ecological success of the eusocial species (2). The collective tasks and functions cannot be performed by any single individual alone. They require a group and inherited behavioral patterns performed by individual workers."

(And later ... [I added the paragraph breaks])

"In advanced eusocial insect colonies, sophisticated innate behaviors establishing sociality are limited to the worker caste, not to queens. This led to our hypothesis that a dedicated developmental program for social behaviors will be found in the pathway determining the differentiation into the worker castes. ...

Dsx proteins are part of the structurally and functionally conserved doublesex and mab-3–related transcription factor family (Dmrt) and are critical for sex-specific differentiation throughout the animal kingdom (39, 40) and sexual behaviors in insects (9, 41). DsxF is required for the worker-characteristic differentiation of the worker ovary, suggesting that DsxF is a component of the worker caste developmental program (19).

The worker bees do not perform sexual behaviors. However, we found that dsx is also expressed in the worker’s brain (42) suggesting that the gene may have another role unrelated to sexual reproduction but related to promoting social living behaviors in the worker caste."

paper link: https://www.science.org/doi/10.1126/sciadv.adp3953