Apis mellifera, honey bee

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Taxonomy

cellular organisms - Eukaryota - Fungi/Metazoa group - Metazoa - Eumetazoa - Bilateria - Coelomata - Protostomia - Panarthropoda - Arthropoda - Mandibulata - Pancrustacea - Hexapoda - Insecta - Dicondylia - Pterygota - Neoptera - Endopterygota - Hymenoptera - Apocrita - Aculeata - Apoidea - Apidae - Apinae - Apini - Apis - Apis mellifera

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Brief facts

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• Distribution

  Apis mellifera is the most commonly domesticated species of honey bees, probably originated in Tropical Africa and spread from there to Northern Europe and East into Asia. Distribution of the species extends from northern Europe to southern Africa, and from the British Isles to the Ural Mountains, western Iran, and the Arabian peninsula. More than 25 subspecies are currently recognized including Apis mellifera scutellata, which after introduction into Brazil apiculture, colonized much of the western hemisphere in less than 50 years thus accomplishing one of most rapid and spectacular biological invasion known.

• Species

  Study of geographic variation in the mtDNA (mitochondrial DNA) of honey bees has revealed four lineages of mtDNA mitotypes: west European (examples: Apis m. mellifera Linnaeus, A. m. iberiensis Engel), east European (examples: A. m. carnica Pollmann, A. m. ligustica Spinola, A. m. caucasia Pollmann,A. m. anatoliaca Maa), African (examples: A. m. capensis Eschscholtz, A. m. intermissa Maa, A. m. lamarckii Cockerell, A. m. litorea Smith, A. m. monticola Smith, A. m. sahariensis Baldenspreger, and A. m. scutellata Lepeletier de Saint Fargeau), and Middle Eastern or Oriental (bees from southeastern Turkey, Lebanon, Jordan, and Israel).

• Social organization

  The honey bees are eusocial insects: they live in colonies that contain one breeding female, or queen; a few thousand males, or drones (the population of drones varies considerably) and a large population (up to and over 100,000) of sterile female worker bees. Example of true mammalian eusociality is described in Curioscope's entry How to hurt a naked mole-rat?.

• Hive temperature

  Bees have behavioral control of temperature and are able to maintain the hive at 32+-0.6°C regardless of the outside temperature. When too cold, bees contract their flight muscles repeatedly without moving their wings; this behavior generates heat.

• Economic value

  The honey bee's primary commercial value is pollination of various fruit and vegetable crops. Other valuable products of honeybees used by humans are honey, beeswax, and propolis. Honeybees are estimated to contribute billions of dollars a year in pollination services alone to the US economy.

• Honeybee endangerment

  Most important honey bee's pathogens that incur great economic damage are the tracheal mite, Acarapis woodi, and the varroa mite, Varroa destructor. Since the 1980s, wild and domestic hives in United States have been decimated by a dual infestations. U.S. honeybee populations declined 50% for last 50 years.

• Honeybee as a model organism

  Honeybee is a model for learning and memory; for studies of allergic disease; for gerontology research, as the long-lived, egg-producing queen is genetically identical to the much shorter-lived female workers; for studies of venom toxicology; and for studies of infectious diseases in dense societies.

• Vision

  Bees, like humans, posess trichromatic color vision, however, their visual spectrum is shifted toward shorter wavelengths, and their visual acuity is about 170 times poorer than that of humans.

• Complexity of tasks

  Honeybee has brain that weighs less than a milligram and consists of fewer than a million neurons. Despite of this, honeybees learn to discriminate visual patterns and to generalize properties of the patterns (such as orientation and symmetry); they are capable of complex associative recall: colors can trigger recall of patterns, and scents can trigger recall of colors; they perform tasks in a context- and time-dependent manner; and they can count to a maximum of four objects presented sequentially or simulteneously.

• Communications

  Honeybees communicate information about important locations (food or nest sites) around the hive through ritualized body movements, called the "waggle dance".
   
  A swarm's decision making relies on sensing a quorum (a sufficient number of scouts) at the one of the nest sites ratherthan sensing a consensus (agreement of dancing scouts) at the swarm cluster. By this hypothesis, a scout bees "vote" for a site by spending time at it.

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Developmental stages (life cycle)

Honeybee passes through 4 distinct life stages: egg, larva, pupa and adult. The process is called complete metamorphosis.

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• egg the queen bee lays a single soft white egg in each cell of the comb; the egg stage usually takes 3 days
  • embryonic queen eggs destined to become queens are laid in a larger cell
  • embryonic drone egg from which drone is going to hatch was not fertilized, thus, drones carry only the chromosomes of the queen
  • embryonic worker
• hatching egg generally hatches into a larva on the fourth day
• larval MeSH larva is a legless grub that resembles a tiny white sausage; the larva is fed a mixture of pollen and nectar called beebread; larval stage takes place during days 4 through 9; larvae undergo several moltings before spinning a cocoon
  • larval worker larval worker is fed a mixture of pollen and nectar called beebread
  • larval drone larval drones are fed royal jelly (a substance secreted from the glands in the heads of worker bees) and develop in a slightly larger cell than worker bees
  • larval queen larval queens are fed royal jelly only
• pupa MeSH on the ninth day the cell is capped with wax and the larva transforms into the pupa; the pupa is a physical transition stage between the amorphous larva and the hairy, winged adult; the pupa does not eat; pupal stage takes 6 days for a future queen, 11 days for a future worker bee, and about 14 days for a drone, to complete
  • pupal worker
  • pupal drone
  • pupal queen
• adult
  • adult worker life span of worker bee is about 20-40 days in summer and 140 days in winter; worker bees are sterile females; during their adult lives worker bees undergo complex and drastic changes in behavior and physiology; the transitions are on only partly genetically predetermined and mostly are governed by environmental factors: they can be delayed, accelerated or even reversed depending on the needs of the hive
    • hive worker
      • cell-tending worker a young honey bee first 2 days after
        hatching who is responsible for cell
        cleaning and capping, and keeping
        brood warm
      • feeding worker 3 to 11 days old honey bee who is
        responsible for feeding larvae and
        queen
      • housekeeping worker 12 to 17 days old bee who is resposible
        for producing wax, building combs, and
        food handling (packing pollen, recieving
        and transporting nectar within the hive
        
      • guardian worker 18 to 21 days old bee who starts with
        ventilating and proceeds with guarding
        the hive's entrance
    • field worker 22 days and older bee who is visiting flowers, collects pollen, nectar, propolis, and water; the onset of foraging coincides with most remarkable transitions in bee's life style: from relatively constant arrythmic activity to diurnality, from homogeneous to highly heterogeneous environments, from darkness to light, from crawling to flight, etc. Forager bees specialize in the collection of either pollen (protein) or nectar (carbohydrates).
  • adult drone life span of drone is about 21-32 days in spring, 90 days in summer; it does not overwinter; drone is a specialized male bee who designed only for mating with the queen after which he dies; all living drones are therefore virgins
  • adult queen life span of queen be can take up to 2 years and depends on amount of sperm which she recieved during her mating period
    • virgin queen newly hatched queen destroys any other unhatched queens, fights to the death any hatched queens, may destroy her mother, and then takes her mating flights
    • mating queen virgin queen flies to a congregation area where hundreds or thousands of unrelated drones await; the drones pursue the queen and several mate with her in flight; of the 90 million sperm deposited by several males in the queen's oviducts, a mixture of about 7 million are stored in a special pouch in her body called the spermatheca; these sperm will be used, a few at a time, during the queen's life to fertilize her eggs; fertilized eggs laid by a queen become female worker bees and new queens; the queen also lays some unfertilized eggs, which produce the drones
    • laying queen the laying queen secretes a pheromone that spreads from body to body among the worker bees and keeps them uninterested in reproduction on their own; the queen lays about 1500 eggs per day

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Body parts

• stinging organs
  • stinger the stinger is composed of two barbed lancets that are connected to the venom sac; by pulling away from a stinging site the bee leaves her stinger, the venom sac and attached muscles in the stung individual's tissues, and subsequently, dies; the stinger continues to throb for 30 to 60 seconds, injecting additional venom and giving off alarm ordors for other bees; up to 100 µg of venom is injected per sting
  • venom sac
    • venom also, apitoxin; the bee venom is composed primarily of proteins; the active components include vasoactive and hemolytic substances such as mellitin, phospholipid A and hyaluronidase as well as small amounts of histamine, dopamine, and norepinephrine responsible for hypotension and tachycardia; 1.3 mg of the venom per kilogram of body weight (1000 stings or about 90 mg for an average adult) is considered to be a lethal dose

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Africanized honey bees

• Some honeybees were taken from Africa (Tanzania) in 1956 to Brazil as part of a breeding program that would allow the European honeybees (EHBs) to adapt to hot South American climate while retaining their high honey production.
• In 1957, 26 swarms of the imported African bees escaped into the surrounding jungles and started to interbreed with local bees. African gene lineage started to replace European patri- and matrilines by several mechanisms, some of which are still poorly understood. Resulted bees began to take on the more aggressive behaviors of the African bees and become known as Africanized Honey Bees (AHBs). AHBs expanded rapidly and at some point contaminated US bee populations in Southern states (Texas around 1990, Arizona and New Mexico by 1993, and California by 1994).
• Many physiological, behavioral, and learning characteristics of AHBs render them survival and dispersal advantages, among them: greater emphasis on pollen collection and faster conversion of pollen into brood; apparent mating advantage of African drones; faster development of African virgin queens and their increased fighting ability, which gives them more opportunities to eliminate European rivals in queen cells; ability of worker bees to fly longer distances; higher reproduction rates that results in exessive swarming (6-12 times per year as opposed to 2-3 times per year); tolerance to harsh temperatures, and an extreme aggressiveness toward perceived intruders.
• Namely because of their heightened excitability and aggressiveness (not because they are more venomous) the AHBs are very dangerous: they are aggravated by minimal disturbances like the vibrations of a lawnmower, light-colored clothing, perfume, and many other stumuli; they pursue their victim relentlessly up to a distance of a quarter of a mile; they attack collectively and in great numbers. Once disturbed, the colony may remain agitated for 24 hours.
• The AHPs are about 10% smaller than EHBs. In contrast with native feral bees they tend to build colonies near populated areas.
• The recommendation for anyone who is attacked by AHBs is to outrun the bees to safety while protecting their eyes and mouth because the bees are attracted to dark moist openings. Diving is not recommended because swimming speed usually is not enough to get away from the bees who also will focus on stinging the head. The bees may even crawl down a tube used for breathing underwater.

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Protocols for studies of learning in the honeybee.

Four main protocols are used for studying learning and memory in honebee.

• Conditioning of the approach flight towards a visual target in free-flying bees.
• Olfactory conditioning of the proboscis extension reflex in harnessed bees.
• Mechanosensory conditioning of the proboscis extension reflex in harnessed bees.
• Olfactory conditioning of the sting extension reflex in harnessed bees.

Appendix I: anatomy of honeybee ovary

Dearden PK. Germ cell development in the Honeybee (Apis mellifera); vasa and nanos expression. BMC Dev Biol. 2006 Feb 17;6:6. PMID: 16503992

Structure of the Honeybee ovary. A) Diagram of the morphology of an ovariole from a mated queen bee ovary. i) the late vitellarium, ii) the early vitellarium, iii) the germarium and terminal filament (the placement of germ cells in the terminal filament is unclear and thus not diagrammed). B) A projection of 25 confocal Z sections through the late germarium of a mated honeybee queen ovariole stained for DNA using Propidium iodide (red) and cortical actin using Alexa fluor 488 phalloidin (green). C) A single confocal section through the germarium of a honeybee queen ovariole, stained as per B, Note the circular actin-rich structures, possibly ring canals. D) Diagram of the morphology of a worker bee ovariole.

Apis mellifera have polytrophic meroistic ovaries, similar to those of Drosophila melanogaster. Despite this similarity, a number of differences in morphology and biology exist. One particularly important difference is that worker bee ovaries have many less ovarioles than those of queens. Workers in a queenright colony have small ovaries that are chemically repressed by the presence of the queen and her eggs. Removal of the queen from a colony can cause the reactivation of the worker bee ovaries and the workers may lay eggs.

Meroistic

In meroistic insect ovaries divisions of gonial cells lead to formation of clusters of sibling cells which remain interconnected by intercellular bridges forming a syncytium. In the meroistic ovary, the oocyte synthesizes little if any RNA. Most of the RNA which accumulates in the oocyte is synthesized by trophocytes.

Polytrophic

In the polytrophic meroistic ovary each ovariole contains a number of clusters of germ cells that form separate structural and functional units termed egg chambers. Each oocyte is associated with a cyst containing 1,3,7 or 15 trophocytes. The trophocytes are derived from the same cell as the oocyte.

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Appendix II: reducing honeybee queen fighting ability

Dietemann V, Zheng HQ, Hepburn C, Hepburn HR, Jin SH, Crewe RM, Radloff SE, Hu FL, Pirk CW. Self assessment in insects: honeybee queens know their own strength. PLoS One. 2008 Jan 9;3(1):e1412. PMID: 18183293

Three A. m. ligustica queens (circled) coexisting peacefully within a colony.

In China, beekeepers found a way to prevent queens from killing one another by ablating their mandibles. By forcing several queens to cohabit, they create more productive colonies for commercial exploitation.

Interestingly, queens with ablated mandibles refrain from engaging in lethal contests that typically characterize their reproductive dominance behavior and coexist peacefully within a colony. This suggests that weak queens exploit an alternative reproductive strategy and provides an explanation for rare occurrences of queen cohabitation in nature. The results of experiments described in theis article further indicate that self-assessment, but not mutual assessment of fighting ability occurs prior to and during the agonistic encounters.

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Appendix III: Asiatic honeybees understand European honeybees

Su S, Cai F, Si A, Zhang S, Tautz J, Chen S. East learns from West: Asiatic honeybees can understand dance language of European honeybees. PLoS One. 2008 Jun 4;3(6):e2365. PMID: 18523550

Apis cerana cerana (Acc) bees (dark abdomens) following the dance of a marked and an unmarked Apis mellifera ligustica (Aml) forager (lighter abdomens) in the mixed-species colony. Both dancers had been trained to an artificial feeder 200 m away from the hive.

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If you see message "Windows Media Player cannot find the file.", please download the file to your computer to play it or go to the original article and play the movie from there. For more movies please see the original article (Open Access).

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Appendix IV: amTOR knockdown blocks queen fate and results in workers

Patel A, Fondrk MK, Kaftanoglu O, Emore C, Hunt G, Frederick K, Amdam GV. The making of a queen: TOR pathway is a key player in diphenic caste development. PLoS One. 2007 Jun 6;2(6):e509. PMID: 17551589

Honey bees (Apis mellifera) provide a principal example of diphenic development. Excess feeding of female larvae results in queens (large reproductives). Moderate diet yields workers (small helpers). The signaling pathway that links provisioning to female developmental fate is not understood. Authors of the article tested their hypothesis that it could include TOR (target of rapamycin), a nutrient- and energy-sensing kinase that controls organismal growth.

Effect of amTOR suppression on caste characters in honey bees. In comparison to control (GFP), amTOR RNAi (RNAi): a, reduced larval growth (exemplified by 4-day-old larvae in sections of a microtiter plate, larval volume is mean±s. e. arbitrary units a.u.); delayed development, b–c, where b is a snapshot of the full phenotypic variance in each group (individuals were 19 days old) – controls emerged with queen morphology, or had advanced pigmentation (indicates more rapid development) and large size. RNAi bees were lightly pigmented and small in size; the latter effect is shown also in d, the wet-weight at adult emergence and in e, the final adult size. Adult size did not overlap between treatment groups (e vs. f, where f shows the two control bees with lowest wet-weight: 155 and 161 mg; weight-range after RNAi was 108–136 mg). amTOR RNAi also reduced ovary size g, to ovariole numbers characteristic of workers (GFP range was 12–180, RNAi range was 2–7 ovarioles per ovary). In sum, the divergence between control and amTOR RNAi is characteristic of queen vs. worker development. Bars are means±s. e. (three asterisks P<1.0 10⁻³). Scale bars a: 10 mm; b, e, f: 5 mm.

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Appendix V: Honey Bee Colony Losses in the U.S., Fall 2007 to Spring 2008

van Engelsdorp D, Hayes J Jr, Underwood RM, Pettis J. A survey of honey bee colony losses in the U.S., fall 2007 to spring 2008. PLoS ONE. 2008; 3(12): e4071.

Honey bee health is challenged on many fronts. Parasites, such as varroa mites (Varroa destructor), honey bee tracheal mites (Acarapis woodi), fungal, bacterial and viral diseases, and kleptoparasites such as small hive beetles (Aethina tumida), many of which have been introduced over the last 20 years to the continental U.S., are all challenges faced by beekeepers. In 2006, a poorly understood phenomenon, Colony Collapse Disorder (CCD), resulted in widespread losses in the U.S.

Between 0.75 and 1.00 million honey bee colonies are estimated to have died in the United States over the winter of 2007–2008. This article is an extensive survey of U.S. beekeepers across the continent, serving as a reference for comparison with future losses as well as providing guidance to future hypothesis-driven research on the causes of colony mortality.

Total colony loss (%) by state. A common practice in epidemiology is to look for spatial patterns to the occurrence of a disease or syndrome. With honey bee colonies making multiple moves around the country it is difficult to assign a colony loss to one region of the country. Losses were assigned to a specific state by the beekeeper and total losses varied by state with no discernable pattern.

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References

PubMed articles

• Mitchell A. Africanized killer bees: a case study. Crit Care Nurse. 2006 Jun PMID: 16717384
• Danforth B. Bees. Curr Biol. 2007 Mar 6;17(5):R156-61. PMID: 17339010
• Elekonich MM, Roberts SP. Honey bees as a model for understanding mechanisms of life history transitions. Comp Biochem Physiol A Mol Integr Physiol. 2005 Aug PMID: 15925525
• Scott Schneider S et al. The African honey bee: factors contributing to a successful biological invasion. Annu Rev Entomol. 2004 PMID: 14651468
• Abramson CI, Aquino IS. Behavioral studies of learning in the Africanized honey bee (Apis mellifera L.). Brain Behav Evol. 2002 PMID: 12097861
• Srinivasan MV. Honey Bees as a Model for Vision, Perception, and Cognition. Annu Rev Entomol. 2009 Sep 3. PMID: 19728835
• The truth about birds and the bees. Environ Health Perspect. 1998 Sep;106(9):A424-5. PMID: 9841208
• Giurfa M. Behavioral and neural analysis of associative learning in the honeybee: a taste from the magic well. J Comp Physiol A Neuroethol Sens Neural Behav Physiol. 2007 Aug;193(8):801-24. PMID: 17639413
• PubMed: free full text articles about bees

Websites

• honeybee.tamu.edu
• Honeybee Research Home