The tunnels they create are lined with silk, which entangles and starves emerging bees. Destruction of honeycombs also result in honey leaking and being wasted. Temperature treatments are possible, but also distorts wax of the honeycombs. Chemical fumigants, particularly CO 2 , are also used. The only domesticated species of honey bee are A. In Japan, where mellifera is vulnerable to local hornets and disease, the Japanese honey bee a. Modern hives also enable beekeepers to transport bees, moving from field to field as the crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide, revising the historical role of the self-employed beekeeper, and favoring large-scale commercial operations.
Bees of various types other than honey bees are also domesticated and used for pollination or other means around the world, including Tetragonula iridipennis in India, the blue orchard bee for tree nut and fruit pollination in the United States, and a number of species of Bombus bumblebees for pollination in various regions globally, such as tomatoes , which are not effectively pollinated by honey bees. Primarily in places where the bee was imported by humans, periodic collapses in honeybee populations have occurred at least since the late 19th century. This has been dubbed "colony collapse disorder" CCD and was at first unexplained.
Honey is the complex substance made when bees ingest nectar, process it, and store the substance into honey combs. Worker bees of a certain age secrete beeswax from a series of exocrine glands on their abdomens. As with honey, beeswax is gathered by humans for various purposes such as candle making, waterproofing, soap and cosmetics manufacturing, pharmaceuticals, art, furniture polish and more.
Bees collect pollen in their pollen baskets and carry it back to the hive. Worker bees combine pollen, honey and glandular secretions and allow it to ferment in the comb to make bee bread. The fermentation process releases additional nutrients from the pollen and can produce antibiotics and fatty acids which inhibit spoilage. In the hive, pollen is used as a protein source necessary during brood-rearing.
In certain environments, excess pollen can be collected from the hives of A. The product is used as a health supplement. It has been used with moderate success as a source of pollen for hand pollination. Bee brood — the eggs, larvae or pupae of honeybees — is nutritious and seen as a delicacy in countries such as Australia, Indonesia, [46] Mexico, Thailand, and many African countries; it has been consumed since ancient times by the Chinese and Egyptians. Propolis is a resinous mixture collected by honey bees from tree buds, sap flows or other botanical sources, which is used as a sealant for unwanted open spaces in the hive.
Royal jelly is a honey-bee secretion used to nourish the larvae. Honey bees have three castes: Males, or drones, are typically haploid , having only one set of chromosomes , and primarily exist for the purpose of reproduction. Diploid drones may be produced if an egg is fertilized but is homozygous for the sex-determination allele. Drones take 24 days to develop, and may be produced from summer through to autumn, numbering as many as per hive. They do not defend the hive or kill intruders, and do not have a stinger. Workers have two sets of chromosomes. Workers typically develop in 21 days.
Beekeeping
A typical colony may contain as many as 60, worker bees. Their duties change upon the age of the bee in the following order beginning with cleaning out their own cell after eating through their capped brood cell: Workers have morphological specializations, including the pollen basket corbicula , [61] abdominal glands that produce beeswax, brood-feeding glands, and barbs on the sting.
Under certain conditions for example, if the colony becomes queenless , a worker may develop ovaries. Worker honey bees perform different behavioural tasks that cause them to be exposed to different local environments. Queen honey bees are created when worker bees feed a single female larvae an exclusive diet of a food called " royal jelly ". In addition to the greater size of the queen, she has a functional set of ovaries, and a spermatheca, which stores and maintains sperm after she has mated.
Apis queens practice polyandry , with one female mating with multiple males. The highest documented mating frequency for an Apis queen is in Apis nigrocincta , where queens mate with an extremely high number of males with observed numbers of different matings ranging from 42 to 69 drones per queen. Once mated, queens may lay up to 2, eggs per day. When a fertile female worker produces drones, a conflict arises between her interests and those of the queen.
The worker shares half her genes with the drone and one-quarter with her brothers, favouring her offspring over those of the queen. The queen shares half her genes with her sons and one-quarter with the sons of fertile female workers. This relationship leads to a phenomenon known as "worker policing". In these rare situations, other worker bees in the hive who are genetically more related to the queen's sons than those of the fertile workers will patrol the hive and remove worker-laid eggs.
Another form of worker-based policing is aggression toward fertile females. In very rare instances workers subvert the policing mechanisms of the hive, laying eggs which are removed at a lower rate by other workers; this is known as anarchic syndrome. Anarchic workers can activate their ovaries at a higher rate and contribute a greater proportion of males to the hive.
Although an increase in the number of drones would decrease the overall productivity of the hive, the reproductive fitness of the drones' mother would increase. Anarchic syndrome is an example of selection working in opposite directions at the individual and group levels for the stability of the hive. Under ordinary circumstances the death or removal of a queen increases reproduction in workers, and a significant proportion of workers will have active ovaries in the absence of a queen.
The workers of the hive produce a last batch of drones before the hive eventually collapses. Although during this period worker policing is usually absent, in certain groups of bees it continues. According to the strategy of kin selection , worker policing is not favored if a queen does not mate multiple times. Workers would be related by three-quarters of their genes, and the difference in relationship between sons of the queen and those of the other workers would decrease.
The benefit of policing is negated, and policing is less favored. Experiments confirming this hypothesis have shown a correlation between higher mating rates and increased rates of worker policing in many species of social hymenoptera. All honey bees live in colonies where the workers sting intruders as a form of defense, and alarmed bees release a pheromone that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species and virtually all other Hymenoptera by the possession of small barbs on the sting, but these barbs are found only in the worker bees.
The sting apparatus, including the barbs, may have evolved specifically in response to predation by vertebrates, as the barbs do not usually function and the sting apparatus does not detach unless the sting is embedded in fleshy tissue. While the sting can also penetrate the membranes between joints in the exoskeleton of other insects and is used in fights between queens , in the case of Apis cerana japonica , defense against larger insects such as predatory wasps e. Asian giant hornet is usually performed by surrounding the intruder with a mass of defending worker bees, which vibrate their muscles vigorously to raise the temperature of the intruder to a lethal level "balling".
Defense can vary based on the habitat of the bee. In the case of those honey bee species with open combs e. Another act of defense against nest invaders, particularly wasps, is "body shaking," a violent and pendulum like swaying of the abdomen, performed by worker bees. The sting and associated venom sac of honey bees are modified so as to pull free of the body once lodged autotomy , and the sting apparatus has its own musculature and ganglion , which allows it to keep delivering venom once detached.
The embedded stinger continues to emit additional alarm pheromone after it has torn loose; other defensive workers are thereby attracted to the sting site. The worker dies after the sting becomes lodged and is subsequently torn loose from the bee's abdomen. The honey bee's venom, known as apitoxin , carries several active components, the most abundant of which is melittin , [77] and the most biologically active are enzymes , particularly phospholipase A2.
Bee venom is under laboratory and clinical research for its potential properties and uses in reducing risks for adverse events from bee venom therapy , [79] rheumatoid arthritis , [80] and use as an immunotherapy for protection against allergies from insect stings. With an increased number of honey bees in a specific area due to beekeeping, domesticated bees and native wild bees often have to compete for the limited habitat and food sources available.
To resolve the issue and maximize both their total consumption during foraging, bumblebees forage early in the morning, while honey bees forage during the afternoon. Honey bees are known to communicate through many different chemicals and odors, as is common in insects. They also rely on a sophisticated dance language that conveys information about the distance and direction to a specific location typically a nutritional source, e.
The dance language is also used during the process of reproductive fission, or swarming, when scouts communicate the location and quality of nesting sites. The details of the signalling being used vary from species to species; for example, the two smallest species, Apis andreniformis and A. Apis mellifera carnica honey bees use their antennae asymmetrically for social interactions with a strong lateral preference to use their right antennae.
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There has been speculation as to honeybee consciousness. The bee was used as a symbol of government by Emperor Napoleon I of France. The priestess at Delphi was the "Delphic Bee". The Quran has a chapter titled " The Bee ". In ancient Egyptian mythology, honeybees were believed to be born from the tears of the Sun God , Ra. A community of honey bees has often been employed by political theorists as a model of human society, from Aristotle and Plato to Virgil. The state of Utah is called the "Beehive State", the state emblem is the beehive, the state insect is the honey bee, and a beehive and the word "industry" appear on both the state flag and seal.
Honey bee life cycle - Wikipedia
A coloured dot applied by a beekeeper identifies the queen. Eastern honey bee A. From Wikipedia, the free encyclopedia. For other uses, see Honey bee disambiguation. Pollination management and List of crop plants pollinated by bees. This section needs additional citations for verification.
Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. June Learn how and when to remove this template message. Bee learning and communication. Giant honey bee A. Journal of Hymenoptera Research. Anatomy of the Honey Bee. Retrieved February 26, Retrieved February 21, Archived from the original PDF on February 29, Proceedings of the California Academy of Sciences. Apini inferred from nuclear and mitochondrial DNA sequence data".
Molecular Phylogenetics and Evolution. Apini inferred from nuclear and mitochondrial DNA sequence data " ". Apidae in South Sulawesi, Indonesia". Journal of the Kansas Entomological Society.
Introduction
Gould; Carol Grant Gould Archived from the original on 4 March Retrieved 12 March Cambridge, Massachusetts and London, England: In some places the researchers considered, wild insects were pollinating most of the plants despite rented honey bees being present. Honeybees also spread exotic plants and transmit pathogens, both of which have been shown to harm other pollinators. Quote checkers like this one, and this one agree. But debunking its message? For another, the statement is hyperbolic and wrong and Einstein was rarely wrong.
Behavioral Ecology and Sociobiology. The Vanishing Honeybee Apis Mellifera However, as humans continued to manipulate the honeybee and deliberately transferred them on a global scale, diseases simultaneously spread and harmed managed colonies. Colony losses have occurred periodically throughout history. Fungus, mites, and starvation have all been thought to be the cause of the deaths. Journal of Invertebrate Pathology.
Retrieved 21 October Retrieved 2 July Archived from the original on 6 February Retrieved 23 October Archived from the original on Retrieved 14 February Retrieved 15 May Home technologies and practices for small agricultural producers, UN Food and Agriculture Organization. The natural history and significance of resin use by honey bees". National Library of Medicine. ID ; "fish protein" ID ; acidic water-based, non-alcoholic flavoured beverages containing calcium in the range of 0.
FDA warned company about making medical claims for bee-derived products". Food and Drug Administration. Visual Dictionary, QA International. Retrieved 18 May Journal of Evolutionary Biology. Behavior Ecology and Sociobiology, Worker policing persists in a hopelessly queenless honey bee colony Apis mellifera. An Introduction to Behavioral Ecology. In temperate regions, worker life history depends strongly on season. The nurse—forager transition is responsive to changing social conditions and can be accelerated, delayed, or even reversed depending on the demographic composition of the hive Robinson ; Huang and Robinson Caste differentiation for larvae that become queens or workers has been associated with the quantity and quality of food that larvae receive before the third larval instar.
Larvae fed a rich diet composed of royal jelly develop into queens, whereas larvae fed a diet composed of glandular secretions, honey, and pollen develop into workers Winston Physiologically, queen-destined larvae have high levels of juvenile hormone JH , which prevents ovarian apoptosis in 5th-instar queen larvae. JH levels have also been implicated in upregulating ecdysteroid titers, which, in turn, activate transcription of genes that direct differentiation to the queen pathway Page and Peng , and references therein.
The insulin signaling and TOR target of rapamycin pathways act as central regulators in queen—worker differentiation Wheeler et al. The nurse—forager transition appears to be orchestrated by JH, which exhibits an age-related increase in worker bees: Winter bees possess well developed hypopharyngeal glands, high vitellogenin levels, and low levels of JH Fluri et al. Social insect queens, including queen honey bees, are extraordinarily long-lived compared both to non-social insects and workers of their own species. The combination of advanced sociality, long life and high fecundity is a syndrome that has evolved independently multiple times in insects, including termites, ants, and several species of bees.
Although difficult to test experimentally, a plausible theory for the evolution of this syndrome has been proposed Keller and Genoud A well-developed population-genetic model of the evolution of aging suggests that most multicellular organisms senesce because, while natural selection strongly opposes alleles that have detrimental effects on early life survival and fecundity, selection is very weak against alleles with equivalent effects that are confined to late life.
Under this general model, either purely detrimental mutations or pleiotropic mutations with both beneficial and detrimental effects can contribute to the evolution of senescence, if the detrimental effects are confined to late ages. These two versions of the model are referred to as the Mutation Accumulation and Antagonistic Pleiotropy models, respectively see Hughes and Reynolds and references therein for more detailed discussion of these models.
Under either version of the general evolutionary model, two demographic patterns can compensate for the decline in selection with age and hence promote long lifespan: Social insect colonies meet both these criteria. Adult queens are protected from predators and environmental extremes by the physical structure of their nests and defensive behaviors of non-reproductive workers. Another critical feature demonstrating the value of social insects as models for aging research is that the trade-off between longevity and reproductive output characterizing other taxa appears to be absent in social insects.
In non-eusocial species of animals, these two traits are typically negatively correlated. Indeed, the existence of such tradeoffs is a cornerstone of life-history theory Roff ; Stearns However, consideration of the ecology of social insects suggests a solution to this puzzle. Life-history tradeoffs are thought to arise from the need to partition limited resources among the competing demands of growth, survival, and reproduction. Because of their colonial life-style and division of labor, eusocial insects societies are thought to be able to garner more energy for the queen than a female of a non-social species would be able to collect Wilson ; Keller If so, then life-history tradeoffs probably do occur in these species, but they are not apparent in contrast to other taxa because their resource budgets are much higher.
There have been few challenges to the general evolutionary model of social insect longevity although the relative contributions of Mutation Accumulation and Antagonistic Pleiotropy processes have been debated, see Hughes and Reynolds and references therein. However, explanations for lifespan differences between queen and worker honey bees have received limited attention.
One plausible hypothesis is that workers, once they become foragers and leave the protected hive environment, experience higher extrinsic mortality due to predation and accident Neukirch ; Schmid-Hempel and Wolf ; Visscher and Dukas An alternate hypothesis is that queens and workers exhibit large differences in intrinsic patterns of aging.
This is the view that motivates the use of comparisons between queens and workers as a tool for understanding the basic biology of aging Parker et al. It is surprising, therefore, that there was little direct evidence to support differences in intrinsic patterns of aging until recently. Early studies of honey bee aging established that worker bees emerging in the spring or fall lived longer than those emerging in the summer Free and Spencer-Booth ; Fukuda and Sekiguchi A few studies pointed to intrinsic senescence as a cause of the worker-queen lifespan difference, but others implicated extrinsic mortality.
Ribbands reported that bees that begin to forage later in life have shorter foraging careers and attributed this pattern to intrinsic senescence. Measurements of glycogen levels in free-living bees led Neukirch to propose that older foraging workers undergo a physiological change that directly affects life expectancy. Neukirch suggested that young foragers can replenish glycogen reserves, but that this ability is impaired in older foragers that exhaust their energy supplies and die in the field. Using experimental manipulation of foraging opportunity, Schmid-Hempel and Wolf found that workers had fixed lifespans regardless of time spent outside the hive, and concluded that lifespan did not depend on extrinsic mortality.
In a study in which workers were trained to visit an artificial flower to collect food, and thus protected from extrinsic mortality, old foragers took longer to collect food and fly between the hive and artificial flower, again suggesting that foragers experienced a decline in flight performance due to senescence Tolfiski Moreover, mortality rate was constant with age in these bees. These observations led the authors to conclude that lifespan in these bees was limited by predation or other sources of extrinsic mortality. In an explicit effort to compare intrinsic vs extrinsic mortality factors in the determination of worker lifespan, and using larger sample sizes than previous studies, Rueppell et al.
They also varied the amount of time bees were allowed to forage in a flight cage. Free-foraging mortality was higher than flight-cage mortality, but both groups showed increasing mortality rates with age consistent with senescence.
Limiting foraging opportunities in the cage had no overall effect on lifespan. There was also a negative correlation between age at first foraging and foraging lifespan, suggestive of pre-foraging senescence. In a separate study, Rueppell et al. They found that workers experienced an increase in mortality with chronological age, but their performance in behavioral assays related to foraging activity did not decline with age. To eliminate the confounding effects of increased extrinsic hazard and energy expenditure faced by foraging bees, Remolina et al.
They tested whether older nurses were more susceptible to different kinds of stress: They therefore concluded that intrinsic senescence affects nurse lifespan independently of extrinsic mortality force. Although there are a few contradictory reports, overall the evidence supports the proposition that worker bees both nurses and foragers exhibit senescence. However, no study has directly assessed queen-worker differences in aging rates. These results suggest that, while workers are experiencing senescent decline in stress resistance by this age, and are not expected to live much longer, queens are highly stress resistant and expected to live more than fold longer.
A reasonable interpretation of this result is that at least part of the difference in queen and worker life span can be attributed to differences in intrinsic rates of aging. Nonetheless, a direct comparison of queen and worker lifespan, and assessment of stress resistance throughout the lifespan, would place the intrinsic senescence hypothesis on firmer footing. Direct comparisons of queen and worker lifespan, and other aspects of age-related performance will be difficult, at least in honey bees.
In this species, it is problematic to house queens and workers under equivalent conditions. Workers cannot be confined for prolonged periods because they must fly in order to defecate. Queens cannot be kept in isolation from workers, since they do not feed themselves. Queens are fed and groomed by nurse bees, making it difficult to house them in the laboratory for long periods.
It might be possible to make rearing conditions more equivalent by, for example, restricting the amount of time workers are allowed to fly. However, energy expenditure, and other differences in lifestyle between queens and workers probably cannot be made entirely equivalent.
As outlined above, there is a well-accepted model for the evolution of long life in social insect queens. However, how queens achieve both long life and sustained high fecundity, compared to other insects, is not yet understood. Several proximate mechanisms for this phenomenon have been proposed; here we review three of the most prominent: The oxidative stress theory of aging Harman states that cumulative oxidative damage causes aging and that lifespan is inversely related to the rate at which damage occurs. Therefore, a long-lived organism, such as the queen honey bee, should have a lower rate of reactive oxygen species ROS production, eliminate ROS more effectively, or better repair oxidative damage in order to minimize accumulation of damage.
The first direct test of this hypothesis in a social insect was conducted by Parker et al. Using honey bees, Corona et al. Confirming the results reported by Parker et al. These two studies indicate that increased antioxidant expression is not a prerequisite for long lifespan in social insects, but do not rule out the possibility that increased lifespan is related to lower ROS production, an alternate mechanism of ROS scavenging Corona et al.
Another hypothesis suggested by Haddad et al. It is possible that honey bees use less well studied molecules for ROS scavenging. Oxidative stress resistance in workers has been linked to the expression of vitellogenin Vg Seehuus et al. Vitellogenin is a female-specific glycolipoprotein synthesized in the fat body of the abdomen, and transported to the ovaries and other body tissues Amdam et al.
Consistent with these findings, Corona et al. Insects possess cellular and humoral immune mechanisms that identify and eliminate foreign tissue and pathogens. Cell-mediated immunity involves hemocytes, which are able to recognize and phagocytose foreign bodies present in the hemolymph. Hemocytes also have the ability to form groups that encapsulate large foreign bodies and nodules to trap invading bacteria.
In addition to cell-mediated immunity, insects synthesize antibacterial peptides in the fat body in response to microbial infection Klowden Moreover, honey bees and other social insects employ behavioral strategies, such as grooming, to fight pathogens reviewed in Evans A general feature of aging in animals is the deterioration of immune efficiency, or immunosenescence Solana and Pawelec A decline with age in the encapsulation and melanization wound response has been observed in worker bumble bees with no corresponding decrease in hemocyte numbers Doums et al.
That conclusion was based on a social manipulation that forced foragers to reverse to the nursing stage. This manipulation resulted in reverted old nurses that were physiologically similar to normal-age nurses low JH titers, high Vg titers, enlarged hypopharyngeal glands and had higher hemocyte counts than foragers of the same chronological age.
In contrast, Schmid et al.
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Neither of these studies linked measures of immune system components directly to mortality rates or senescence, and both contrast with a report that challenges to the immune system of bumble bees, which should increase immune component expression, actually led to decreased survival under starvation conditions Moret and Schmid-Hempel In fact, chronic immune system activation can cause decreased lifespan in the fruit fly Drosophila melanogaster Libert Thus, it is not clear that upregulation of immune system components, should be directly associated with longer lifespan.
The insulin insulin-like signaling IIS pathway regulates metabolism, development, reproduction, and longevity in model organisms, including nematode worms, fruit flies, and, and mammals reviewed in Kenyon ; Partridge and Gems , Hughes and Reynolds ; Sinclair et al. In addition to its potential effect on caste determination during development Wheeler et al.
Recent studies suggest that the IIS pathway interacts with Vg and JH to regulate honey bee lifespan, and these interactions may provide the key to both the long life and high fecundity of the queen Corona et al. Similar models of endocrine regulation have been proposed to explain life history variation between nurses and foragers and between workers in divergent selection lines Amdam and Omholt b ; Amdam et al.
Here we focus on endocrine signaling differences that may lead to variation in lifespan between queen and worker honey bees. In Drosophila, JH and Vg abundance are positively correlated, and a calorically enriched diet causes increased signaling through the IIS pathway, which in turn shortens lifespan Tu et al.