Honey bees (Apis mellifera) rely on a
complex diet to maintain colony health, productivity, and resilience. Among the
most vital components of their nutrition are proteins and carbohydrates. These
macronutrients not only fuel daily activity but also influence brood
development, immunity, longevity, and foraging efficiency. Understanding their
roles and dietary sources is essential for both traditional and modern
beekeeping, especially as environmental changes and habitat loss continue to
affect floral diversity.
The Role of Proteins
in Honey Bee Physiology
Proteins are critical for the growth
and development of honey bee larvae, as well as for the maintenance and repair
of tissues in adult bees. They are especially important in the production of
royal jelly, brood food, enzymes, and immune proteins.
Nurse bees require high protein intake
for gland development, particularly the hypopharyngeal glands, which secrete
larval food (Crailsheim et al., 1992). Protein deficiency can lead to reduced
brood rearing and impaired queen development (DeGrandi-Hoffman et al., 2010).
Pollen is the primary protein source
for bees. Its protein content can vary from 2% to over 60%, depending on plant
species (Roulston et al., 2000). Bees instinctively prefer pollen from species
that offer not only high protein but also a balanced amino acid profile (Cook
et al., 2003).
The Role of
Carbohydrates in Honey Bee Energy Metabolism
Carbohydrates serve as the principal
energy source for bees. They power all energy-demanding activities including
foraging, thermoregulation, wax secretion, flight, and communication dances.
Nectar, which is primarily composed of
sucrose, glucose, and fructose, provides bees with their carbohydrate
requirements. Adult worker bees convert nectar into honey, which they store for
future energy needs (Brodschneider & Crailsheim, 2010). During periods of
dearth or winter, honey becomes the critical survival resource for the colony.
A deficiency in carbohydrates leads to
low energy reserves, reducing the colony’s ability to thermoregulate or forage
efficiently. In extreme cases, it may result in colony starvation even when
protein is adequate.
The diversity of flora around the
apiary is essential. Monofloral pollen diets are nutritionally inferior
compared to polyfloral ones (Alaux et al., 2010). Access to multiple plant
species ensures a balanced nutrient intake.
Nutritional
Imbalance and Supplementation
Due to intensive agriculture and loss
of native forage, bees often face nutritional stress. Beekeepers can intervene
by providing protein supplements such as pollen substitutes or patties made
from soy flour, brewer’s yeast, or milk powder. These are particularly helpful
during early buildup or pollen dearths (Herbert, 1992).
Sugar syrup is commonly used to
supplement carbohydrates, especially before winter or during nectar scarcity.
However, over-reliance on artificial feeds without monitoring colony health can
disrupt microbial balance and immunity (Standifer, 1980).
Nutritional
Influence on Colony Health
Proper protein and carbohydrate
nutrition has been linked to:
Enhanced disease resistance (Alaux et al., 2010)
Higher queen and brood viability (DeGrandi-Hoffman et al., 2010)
Honey bees communicate extensively through pheromones, with most research focused on the queen and workers. Drones (male bees) also produce pheromones, although far less is
known about them. Recent studies have begun to illuminate the chemical signals male honey bees produce, when they produce them, and how these cues function in mating and colony dynamics (Gryboś et al., 2025).
Pheromone Sources
and Chemical Nature in Drones
Male honey bee pheromones derive
primarily from the drones’ mandibular glands. Chemical analyses show that drone
mandibular gland secretions are dominated by fatty acids (both saturated and
unsaturated, some with methyl branching) ranging roughly from 9 to 22 carbons
in length (Villar et al., 2018). Notably, two major components identified are
hexadecanoic acid (palmitic acid) and (Z)-9-octadecenoic acid (oleic acid)
(Lensky et al., 1985). These compounds and a few others make up the bulk of the
drone’s pheromonal blend. In contrast to queens – whose mandibular glands
produce potent mixtures like 9-ODA and other specialized acids – the drones’
blend consists mostly of relatively simple fatty acids. Drones also lack
certain pheromone glands present in workers; for example, males have no sting
(hence no alarm pheromone) and do not use Nasonov orientation pheromone,
reflecting their limited roles in colony labor. While small tarsal (foot)
glands exist in males, their secretions appear minimal and are not known to
play a significant communicative role in the colony. In short, the drone’s
known pheromones are few, with the mandibular gland being the key source
identified so far.
Onset of Pheromone
Production and Maturation
Drone honey bees do not produce
effective pheromones immediately upon emergence. Pheromone production ramps up
as the drone matures sexually. Drones typically reach sexual maturity about 1–2
weeks after emergence (around 9–12 days old) (Bastin et al., 2017).
Correspondingly, studies show that only sexually mature drones produce the
pheromonal cues that attract others. In laboratory experiments, young drones
(2–3 or 7–8 days old) showed no attraction to the odor of their peers, whereas
sexually mature drones (12–15 days old) were strongly attracted to the scent of
other mature males (Bastin et al., 2017). This indicates that pheromone
production (or release) begins in earnest as drones become capable of mating.
Histological observations confirm that the drone’s mandibular glands develop
and fill with secretions in the first week or so of adult life, then begin to
degenerate around the onset of sexual maturity (Lensky et al., 1985).
Functions in Mating
and Drone Congregation Behavior
Unlike worker or queen pheromones
which regulate complex social tasks in the hive, drone pheromones serve a more
singular purpose: reproduction. The primary function of drone pheromones is to
mediate interactions during mating. In the afternoons of the mating season,
mature drones leave their hives and gather in the air at specific locations
known as drone congregation areas (DCAs) – sometimes thousands of males from
many colonies converge in one spot (Bastin et al., 2017). How do all these
males find the same location? A combination of environmental cues and
pheromones is at work. Landmark cues (such as treelines or topography) help
orient drones to the general area, but in the final localization,
drone-produced pheromones draw males together into the tight congregation
(Wanner & Nishida, 2017).
Decades ago, researchers hypothesized
that an aggregation pheromone emitted by drones facilitates DCA formation
(Lensky et al., 1985). This was first supported when extracts of drone
mandibular glands were found to attract flying drones in field tests (Lensky et
al., 1985). More recently, controlled lab studies confirmed that drones are
indeed attracted to the odor of other drones, validating the existence of a
drone aggregation signal (Wanner & Nishida, 2017). Importantly, this
pheromonal attraction is specific to drones – a synthetic blend of the six main
mandibular fatty acids or the natural gland extract will lure other drones, but
will not attract worker bees inside or outside the hive (Villar et al., 2018).
In other words, the drone pheromone is a releaser signal aimed at fellow males
(and perhaps queens), not a colony-wide messenger.
The ultimate purpose of drone
congregation is to mate with virgin queens. Here, drone pheromones intersect
with queen pheromones. Virgin queen honey bees generally arrive at DCAs after
the drones have already assembled (Bastin et al., 2017). Drones detect and
chase the queen largely by sensing the queen’s own sex pheromone – the queen
mandibular pheromone (QMP), especially its key component 9-oxo-2-decenoic acid
(9-ODA) (Gryboś et al., 2025). In fact, drones have highly tuned antennae and
brain olfactory centers (macroglomeruli) specifically devoted to detecting
9-ODA, which triggers their frenzied pursuit of a queen in flight (Brockmann et
al., 2006). However, queens may also use drone pheromones to their advantage:
lab experiments have demonstrated that virgin queens are attracted to the odor
bouquet of a group of drones (Wanner & Nishida, 2017). A sexually receptive
queen oriented toward an array of drone scent in a wind chamber, whereas she
ignored worker bee odors. This suggests that a drone-produced odor cue helps
queens locate congregation sites in the final stage of their nuptial flight.
Notably, drone pheromones seem to have
little or no direct role in daily colony social dynamics beyond mating. Worker
bees do not exhibit obvious behavioral or physiological responses to the
presence of drone pheromone inside the hive (Villar et al., 2018). Drones are
typically tolerated in the hive during the breeding season due to hormonal and
resource cues rather than any known drone-emitted primer pheromone. In lean
times, workers evict drones in autumn, again seemingly without a specific drone
chemical signal driving the process. Thus, the drone’s chemical communication
is narrowly focused on its reproductive mission, unlike the queen’s pheromones
which influence everything from worker task allocation to suppression of new
queens, or the workers’ pheromones that coordinate foraging, alarm, and brood
care.
Evolutionary and
Ecological Context
The specialized nature of drone
pheromones reflects the drone’s singular role in the honey bee superorganism.
Drones exist only to spread the colony’s genes via mating, and accordingly
their pheromones have evolved for mate location and competition. In many
insects, either the female produces a sex attractant or the males produce
aggregation signals – honey bees have elements of both strategies. The queen’s
powerful sex pheromone (9-ODA in QMP) has an immediate effect of arousing
copulatory behavior in drones and drawing them in toward her (Brockmann et al.,
2006). Drones, on the other hand, appear to cooperate (unwittingly) by emitting
an odor that helps form large congregations, which in turn increases each
male’s chance to encounter a queen (Wanner & Nishida, 2017).
From an evolutionary perspective, a
drone aggregation pheromone likely benefits all participating males by creating
a predictable “mating marketplace” that virgin queens can find. Drones from
many hives mix at DCAs, promoting outbreeding and genetic diversity in the
species. The consistency of DCA locations year after year – some spots have
been used by drones for decades – implies that aggregation cues are robust and
long-standing in honey bee reproductive ecology. Environmental factors (like
terrain landmarks and even geomagnetic anomalies) help drones roughly orient to
a site, but the final assembly into a dense drone cloud likely requires the
pheromone signal as a short-range cue to cluster within a defined space.
Compared to the multi-functional
pheromones of queens and workers, drone pheromones are limited but crucial.
They do not regulate colony labor or development, yet they are indispensable
for successful mating. The existence of a male-produced pheromone in Apis
mellifera also invites comparisons to other bees: for instance, males of
some bumblebee species produce pheromonal scents to mark perch sites and
attract queens, and males of solitary bees often emit pheromones during mating
swarms. Honey bee drones fit into this broader context of male chemical
signaling, but their pheromones had remained elusive until recently. Now, with
advanced analytical chemistry and creative bioassays (like drone walking
simulators), scientists have confirmed that drones indeed contribute their own
chemical voice to the honey bee communication repertoire.
References
Bastin, F., Savarit, F., Lafon, G.,
& Sandoz, J.-C. (2017). Age-specific olfactory attraction between Western
honey bee drones (Apis mellifera) and its chemical basis. PLOS ONE,
12(10), e0185949. https://doi.org/10.1371/journal.pone.0185949
Brockmann, A., Dietz, D., Spaethe, J.,
& Tautz, J. (2006). Beyond 9-ODA: Sex pheromone communication in the honey
bee. Apidologie, 37(2), 138–154. https://doi.org/10.1051/apido:2005076
Gryboś, A., Staniszewska, P., Bryś, M.
S., & Strachecka, A. (2025). The pheromone landscape of Apis mellifera:
Caste-determined chemical signals and their influence on social dynamics. Molecules,
30(11), 2369. https://doi.org/10.3390/molecules30112369
Lensky, Y., Cassier, P., Notkin, M.,
Delorme-Joulie, C., & Levinsohn, M. (1985). Pheromonal activity and fine
structure of the mandibular glands of honeybee drones (Apis mellifera
L.). Journal of Insect Physiology, 31(4), 265–276.
https://doi.org/10.1016/0022-1910(85)90031-3
Villar, G., Wolfson, M. D., Hefetz,
A., & Grozinger, C. M. (2018). Evaluating the role of drone-produced
chemical signals in mediating social interactions in honey bees (Apis
mellifera). Journal of Chemical Ecology, 44(1), 1–8.
https://doi.org/10.1007/s10886-017-0910-2
Wanner, K. W., & Nishida, R.
(2017). Virgin queen attraction toward males in honey bees. Scientific
Reports, 7, 40697. https://doi.org/10.1038/s41598-017-06241-9
Understanding Burr
Comb and Cross Comb in Langstroth Hives
Burr Comb
Burr comb refers to small, irregular
wax structures built by bees in unintended gaps within the hive, such as
between frames, under the lid, or across the inner cover. These structures
arise when spacing within the hive exceeds the ideal bee space, which is
approximately 6 to 9 millimetres. In these situations, bees instinctively fill
the space with wax or propolis (Bradbear, 2009; Seeley, 2010). Langstroth’s
original design emphasized this precise spacing, noting that deviations would
prompt bees to either block tight gaps with propolis or fill wide spaces with
comb (Langstroth, 1853).
Cross Comb
Cross comb occurs when bees construct
comb at angles that do not align with the frames. Instead of building straight
within the plane of a frame, the comb stretches across adjacent frames. This
results in disorganized construction that complicates inspections and often
leads to the destruction of comb
during hive management (Winston, 1987).
Causes of Burr Comb
and Cross Comb
Several factors contribute to the
formation of burr and cross comb. First, improper bee space is a major culprit.
If the space between hive components deviates from the 6–9 mm standard, bees
will likely modify the space by constructing wax bridges or barriers.
Misaligned or warped frames also create uneven gaps, encouraging the bees to
build in unintended directions (Root, 1921).
Additionally, the absence of
foundation or poorly attached foundation sheets can prompt bees to follow their
instincts and create naturally oriented, and often irregular, combs (Hepburn
& Radloff, 2011). Another significant cause is the orientation and leveling
of the hive. Bees rely on gravity to construct vertical combs. If the hive is
tilted, especially from side to side, the bees’ natural alignment is disrupted,
leading to angled combs that result in cross combing (Seeley, 2010).
Furthermore, introducing swarms into
boxes without drawn comb or foundation often results in haphazard comb
building, particularly in deep boxes with no visual or structural guides
(Crane, 1990).
Disadvantages of
Burr Comb and Cross Comb
The disadvantages of burr and cross
comb are numerous. They impede routine inspections by fusing frames together or
obscuring frame edges, making it difficult to lift them without damaging bees
or comb. This increases the risk of injuring the queen or destroying brood and
honey stores. Additionally, broken burr comb filled with honey creates a sticky
mess that attracts pests and disturbs the colony. Over time, irregular comb
reduces usable hive space and complicates colony management (Delaplane, 2007).
Control and
Prevention
To control and prevent burr and cross
comb, several strategies can be implemented. Maintaining precise bee space is
essential. This involves using quality equipment and regularly inspecting for
warped or damaged frames. Providing bees with foundation sheets or comb starter
strips guides proper comb construction, especially in new hives (Hepburn &
Radloff, 2011).
Ensuring that hives are level,
particularly from side to side, supports natural vertical comb alignment.
Frequent early inspections during comb-building phases allow beekeepers to
intervene before cross comb becomes extensive. In addition, using the correct
number of frames to avoid excessive gaps in supers can prevent bees from
constructing comb in unintended spaces.
Corrective actions include trimming
irregular comb early and realigning frames. If done promptly, bees often adapt
and reconstruct comb in the desired direction. Consistent management and
attention to structural detail ensure that Langstroth hives function as
intended, reducing disruptions and supporting colony health.
Raising Queens from Swarm, Supersedure, and Emergency
Cells
In managed beekeeping, the type of queen cell used to raise
a new queen can significantly influence the overall success and productivity of
the resulting colony. Beekeepers commonly rely on swarm, supersedure, or
emergency queen cells when rearing replacement queens or expanding colonies.
However, these cell types vary in terms of biological origin, the circumstances
in which they are produced, and the quality of queens they generate. Drawing
from contemporary scientific literature, this article explores these
differences in depth and outlines when and why each queen cell type might be
preferred.
Queen Cell Types and Their Biological Origins
Swarm cells are produced by colonies in preparation for
swarming—a natural method of colony
reproduction. These elongated, peanut-shaped cells are typically located along
the edges of brood comb and are created under conditions of resource abundance
and strong colony vigor. The queen that lays the eggs destined to become swarm
queens is often in good reproductive condition, and the resulting larvae are
well-fed by nurse bees throughout their development (Simone-Finstrom et al.,
2016). Because they are raised under planned and favorable circumstances, swarm
cells tend to yield robust queens.
Supersedure cells, in contrast, are constructed when the
colony determines that the reigning queen is no longer performing optimally—often due to declining pheromone
output or reduced egg-laying capacity. These cells are usually found in the
central part of the comb. Unlike swarm situations, the colony remains
queenright during supersedure events. The objective here is not reproduction
but internal queen replacement to sustain colony performance. While the
environmental conditions are often stable, the urgency for replacement is
typically lower than in emergency scenarios (Winston, 1987).
Emergency cells arise when a colony loses its queen
abruptly—either through beekeeper error,
predation, or disease. In such cases, the colony responds by identifying young
worker larvae, less than three days old, and rearing them into queens by
feeding them royal jelly and enclosing them in specially extended vertical
cells. This rapid response mechanism ensures the survival of the colony, but it
imposes developmental constraints on the replacement queen due to time
limitations and suboptimal nutritional conditions (Tarpy et al., 2000).
Scientific Insights into Queen Quality
Research indicates that queens reared under swarm
conditions consistently outperform those produced under supersedure or
emergency conditions. Swarm queens tend to be larger in size, exhibit more
developed reproductive organs, and have higher mating success rates. This
superiority is attributed to the favorable colony status and abundant resources
available during their rearing period (Gilley, 2001).
Supersedure queens are generally acceptable, though studies
show a degree of variability in their reproductive fitness depending on the age
and condition of the queen they replace and the overall health of the colony
(Hatch et al., 1999). Emergency queens are more likely to be of lower quality.
Since they are derived from worker-destined larvae and raised hastily, they may
be smaller, less fertile, and more susceptible to mating failures or early
supersedure (Tarpy et al., 2000).
Guidance for Beekeepers: Choosing the Right Cell Type
Swarm cells are best suited for planned colony divisions,
such as raising nucleus colonies (nucs) or executing controlled expansions.
These cells tend to yield queens of excellent quality, but they also require
close monitoring to avoid uncontrolled swarming.
Supersedure cells are most appropriate for replacing a
failing queen within an otherwise stable colony. While these queens are not as
reliably robust as swarm queens, they are often sufficient for colony
maintenance, especially if the workers make the replacement decision gradually
and under stable environmental conditions.
Emergency queen cells should be reserved for true crises—situations in which the colony is
suddenly queenless. While they serve a critical survival function, the
resulting queens are often inferior in terms of reproductive performance and
longevity. In such cases, beekeepers may consider introducing a purchased or
grafted queen instead, to avoid compounding colony stress with a suboptimal
queen.
Considerations Affecting Queen Success
Several additional factors influence the success of queens
reared from any of these cell types. First, timing is crucial. Queens raised
from swarm and supersedure cells develop over the standard 16-day period from
egg to emergence. In contrast, emergency queens often develop from slightly
older larvae, compressing their developmental window and potentially
compromising physical development (Winston, 1987).
Second, nutritional conditions play a fundamental role in
queen quality. Regardless of the cell type, inadequate feeding of larvae during
early development will result in smaller, less capable queens. Even
well-situated swarm cells can produce poor queens if the colony is experiencing
nutritional stress.
Finally, successful mating is essential for a viable queen.
Factors such as weather, availability of drones, and mating flight success all
influence whether a seemingly well-developed queen will establish a strong
laying pattern. A queen’s quality is not solely defined by her physical
characteristics but also by her ability to mate effectively and begin laying
fertilized eggs promptly (Delaney et al., 2009).
References
Delaney, D. A., Keller, J. J., Caren, J. R., & Tarpy,
D. R. (2009). The physical, insemination, and reproductive quality of honey bee
queens (Apis mellifera). Apidologie, 40(5), 563–572. https://doi.org/10.1051/apido/2009049
Gilley, D. C. (2001). The behavior of honey bees (Apis mellifera ligustica)
during swarm cell construction. Animal Behaviour, 61(1), 1–9. https://doi.org/10.1006/anbe.2000.1552
Hatch, S., Tarpy, D. R., & Fletcher, D. J. C. (1999). Worker regulation of
emergency queen rearing in honey bee colonies and the resultant variation in
queen quality. Insectes Sociaux, 46, 372–377. https://doi.org/10.1007/s000400050158
Simone-Finstrom, M., Li-Byarlay, H., Huang, M. H., Strand, M. K., Rueppell, O.,
& Tarpy, D. R. (2016). Migratory management and environmental conditions
affect lifespan and oxidative stress in honey bees. Scientific Reports,
6, 32023. https://doi.org/10.1038/srep32023
Tarpy, D. R., Hatch, S., & Fletcher, D. J. C. (2000). The influence of
queen age and quality during queen replacement in honeybee colonies. Animal
Behaviour, 59(1), 97–101. https://doi.org/10.1006/anbe.1999.1278
Winston, M. L. (1987). The Biology of the Honey Bee. Cambridge, MA:
Harvard University Press.
Before the 1850s, beekeepers commonly used fixed comb hives
such as straw skeps or hollow log gums. In these traditional hives, bees
attached their comb directly to the internal surfaces, making it impossible to
harvest honey or inspect brood without tearing apart the comb (Crane, 1999).
Honey harvesting typically resulted in the destruction of the colony, often by
suffocating the bees using smoke or sulfur (Bradbear, 2009). These limitations
made traditional beekeeping unsustainable and inefficient.
Discovery of the Bee Space (1851)
In 1851, Reverend Lorenzo Lorraine Langstroth, an American
clergyman and beekeeper, observed that when a space of approximately 6 to 9
millimeters was maintained between combs or hive parts, bees neither built comb
in the gap nor sealed it with propolis. He termed this discovery the “bee space” (Langstroth, 1853). Langstroth then
designed a rectangular hive using removable wooden frames suspended inside a
box, with all gaps spaced precisely to preserve the bee space. This innovation
allowed frames to be removed for inspection or honey harvesting without
destroying combs or aggravating the bees (Root, 1918).
Patent of the Movable Frame Hive
(1852)
Langstroth patented his movable frame beehive on October 5,
1852, under U.S. Patent No. 9300. The design incorporated internal spacing that
respected the bee space, allowing beekeepers to manage colonies with
unprecedented ease and efficiency (Graham, 1992). By the end of that year,
Langstroth had operationalized over 100 of these hives in his apiary. The
design made it possible to harvest honey without killing bees or destroying wax
combs, representing a major advancement in the field (Crane, 1999).
Publication and Impact (1853)
In 1853, Langstroth published A Practical Treatise on
the Hive and Honey-Bee, a comprehensive manual explaining his beekeeping
method and hive design (Langstroth, 1853). The book quickly became a
foundational text for American beekeepers, offering detailed guidance on colony
management, seasonal care, and honey extraction. The principles of movable
frames, modular hive construction, and bee space became standard features in
modern hive design.
Global Influence and Legacy
Although European beekeepers like Jan Dzierżon and August von Berlepsch had explored similar concepts
earlier, Langstroth was the first to fully implement the bee space principle in
a practical and replicable hive system (Crane, 1999). His invention transformed
beekeeping from a destructive to a sustainable practice and earned him
recognition as the father of American beekeeping (Root, 1918). The Langstroth
hive quickly spread internationally and remains the most widely used hive
today, forming the basis for approximately 75 percent of the world’s managed hives (Adjare, 1990;
Bradbear, 2009).
References
Adjare, S. O. (1990). Beekeeping in Africa. Food and
Agriculture Organization of the United Nations.
Bradbear, N. (2009). Bees and their role in forest
livelihoods: A guide to the services provided by bees and the sustainable
harvesting, processing and marketing of their products. Food and
Agriculture Organization of the United Nations.
Crane, E. (1999). The world history of beekeeping and
honey hunting. Routledge.
Graham, J. M. (Ed.). (1992). The hive and the honey bee.
Dadant & Sons.
Langstroth, L. L. (1853). A practical treatise on the
hive and honey-bee. Hopkins, Bridgman & Company.
https://www.gutenberg.org/ebooks/24583
Root, A. I. (1918). The ABC and XYZ of bee culture.
A. I. Root Company.
U.S. Patent and Trademark Office. (1852). Movable-frame
beehive: Patent No. 9300. Filed October 5, 1852.
📚 View Full References
Adjare, S. O. (1990). Beekeeping in Africa. FAO.
Bradbear, N. (2009). Bees and their role in forest livelihoods. FAO.
Crane, E. (1999). The world history of beekeeping and honey hunting. Routledge.
Graham, J. M. (Ed.). (1992). The hive and the honey bee. Dadant & Sons.
Langstroth, L. L. (1853). A practical treatise on the hive and honey-bee. Gutenberg. [Link]
Root, A. I. (1918). The ABC and XYZ of bee culture. A. I. Root Co.
U.S. Patent and Trademark Office (1852). Patent No. 9300.
Drone honey bees (Apis mellifera) are
biologically specialized for one function: mating with a queen. Unlike worker
bees, drones do not forage, care for the brood, or build comb. They exist
solely to pass on genetic material. However, this act comes at a fatal cost. A
drone that successfully mates dies immediately afterward. This article explains
the biological mechanism and evolutionary reasoning behind drone mortality
after mating.
Drone Congregation Areas: Where Mating
Begins
Honey bee mating does not occur randomly. Instead, it
takes place at designated locations called Drone Congregation Areas (DCAs).
These are stable open air zones typically found 10 to 40 meters above ground
level. They often form in open spaces near natural features such as forest
edges, tree gaps, or clearings (Koeniger, Koeniger, & Tingek, 2005).
Thousands of drones from different colonies gather
daily in DCAs during mating season. Virgin queens enter these areas during
their nuptial flights and are pursued midair by drones. Mating occurs in
flight, and a queen may visit several DCAs, typically mating with 12 to 20
drones over one or more days (Baudry et al., 1998). This system encourages
genetic mixing and reduces the risk of inbreeding.
The Mating Process and Drone
Physiology
When a drone succeeds in mating, he uses an internal
structure called the endophallus to inseminate the queen. This organ is
everted, or turned inside out, under intense pressure during ejaculation. A
single ejaculation can transfer up to 90 million sperm cells (Koeniger et al.,
2014).
The force of this transfer is so extreme that the
endophallus ruptures and becomes detached inside the queen’s reproductive
tract, forming a temporary mating plug. As a result of this rupture, the drone
experiences fatal abdominal damage, including tearing of tissues and hemolymph
loss. Death follows within seconds (Woyke, 1958).
Why Drones Die After Mating
Drone mortality is not accidental. It is a direct
result of their anatomical specialization. Unlike males in many insect species
who survive and can mate multiple times, honey bee drones are structured for
single use reproduction. The mating event causes irreversible physical damage,
making post mating survival biologically impossible.
Evolutionary Significance
From an evolutionary standpoint, this reproductive
strategy represents a terminal investment. The drone maximizes his reproductive
success through one complete mating. The detachment of the endophallus ensures
full sperm transfer and may briefly deter further mating attempts by other
drones (Baer, 2005).
This seemingly costly strategy
benefits the colony. Queens store the sperm of all mates in a structure called
the spermatheca, using it to fertilize eggs throughout their lives. By mating
with multiple drones, the queen ensures genetic diversity and resilience within
the colony.
References
Baer, B. (2005). Sexual selection in Apis bees. Apidologie,
36(2), 187–200. https://doi.org/10.1051/apido:2005012
Baudry, E., Solignac, M., Garnery, L., Gries, M.,
Cornuet, J. M., & Koeniger, N. (1998). Relatedness among drone
congregations of the honeybee (Apis mellifera L.). Proceedings of the
Royal Society B: Biological Sciences, 265(1392), 2009–2014.
https://doi.org/10.1098/rspb.1998.0533
Koeniger, G., Koeniger, N., & Tingek, S. (2005). Mating
Biology of Honey Bees (Apis). International Bee Research Association.
Koeniger, G., Koeniger, N., Ellis, J., & Connor, L.
J. (2014). Queen Mating and Reproduction in Honey Bee Colonies. Wicwas
Press.
Woyke, J. (1958). Natural and artificial insemination
of queen honeybees. Bee World, 39(3), 57–65.
https://doi.org/10.1080/0005772X.1958.11095037
Sources and Further Reading
You can view or access the sources cited in this article below:
Yes, honey bees (Apis
mellifera) eat the honey they produce. Honey is their primary source of
carbohydrates and is vital not only for daily energy but also for other
metabolic functions such as thermoregulation, brood care, and wax secretion
(Quinlan et al., 2023; Huang, 2018).
Nutritional
Function of Honey
Honey is composed
primarily of simple sugars—fructose and glucose—making up about 80–85% of its content, with water
accounting for 15–17%
(Martinotti & Ranzato, 2023). It also contains trace amounts of enzymes,
amino acids, vitamins, and minerals (Rao et al., 2016). This sugar-rich
composition makes honey an efficient energy source. Worker bees consume honey
to fuel flight, fanning, foraging, and maintaining hive temperature. During
cold periods or nectar scarcity, bees rely entirely on their honey reserves for
survival (Mississippi State University Extension, 2019).
Honey
and Wax Production
Beyond energy, honey
is also essential in wax production. Worker bees consume large amounts
of honey to generate wax scales from abdominal glands. According to Hepburn
(1986), it takes approximately 6.6 to 8.4 kg of honey to produce 1 kg of
beeswax, highlighting the metabolic cost of comb building. The wax is
critical for constructing brood cells and storage areas for pollen and honey.
Honey
Consumption in Tropical Climates
In tropical regions
such as sub-Saharan Africa where winter is absent, honey bees do not enter
dormancy but still rely on stored honey during seasonal nectar dearths. For
example, during long dry spells or periods of intense heat when flowering
plants are sparse, bees reduce foraging and consume stored honey to maintain
basic colony functions (Adjare, 1990; Nuru et al., 2015). These seasonal
shortages mimic winter scarcity in temperate zones in terms of resource stress.
Do
Bees Eat Honey Before Rain or If Harvest Is Delayed?
Yes. Bees can detect
atmospheric changes, including humidity and pressure drops that signal
impending rain. In anticipation, they increase feeding on stored honey to
prepare for confinement during rainy days (Crane, 1990). Similarly, if honey
remains unharvested and environmental conditions change—such as onset of a rainy season—bees may consume surplus honey,
especially if foraging is curtailed. In such cases, beekeepers may find reduced
yields if harvest is delayed.
Furthermore, the timing
of harvesting is crucial. Delayed harvesting during or after peak nectar
flow can coincide with the colony shifting to consumption rather than storage—especially if pollen becomes limited or nectar stops
flowing. A study by Fichtl & Admasu (1994) in Ethiopia noted that local
beekeepers often lose part of the honey yield if not collected early, as bees
consume the surplus during late dry seasons.
How
Much Honey Do Bees Consume?
An adult worker bee
consumes approximately 11 mg of honey daily for metabolic needs (Huang, 2018).
A standard colony of 50,000 bees may consume up to 1 kg of honey per day
under active foraging or brood-rearing conditions. For long-term survival
during scarcity periods, colonies may require 30–45 kg of honey reserves,
depending on region and hive size (Mississippi State University Extension,
2019).
Sources and Downloads
You can view or download the academic and institutional sources cited in this article below:
Adjare, S. O.
(1990). Beekeeping in Africa. FAO Agricultural Services Bulletin 68/6.
Food and Agriculture Organization of the United Nations.
Crane, E. (1990). Bees and Beekeeping: Science, Practice and World Resources.
Heinemann Newnes.
Fichtl, R., & Admasu, A. (1994). Honeybee flora of Ethiopia.
Deutsche Gesellschaft für
Technische Zusammenarbeit (GTZ).
Hepburn, H. R. (1986). Honeybee wax: An overview of its biology, collection,
properties and functions. Bee World, 67(3), 119–132. https://doi.org/10.1080/0005772X.1986.11098954
Huang, Z. (2018). Feeding honey bees (Bulletin E-3369). Michigan State
University Extension.
Martinotti, S., & Ranzato, E. (2023). Applications of beehive products for
wound repair and skin care. Cosmetics, 10(5), 127.
https://doi.org/10.3390/cosmetics10050127
Mississippi State University Extension. (2019). Colony growth and seasonal
management of honey bees.
https://extension.msstate.edu/sites/default/files/publications/publications/P3052_web.pdf
Quinlan, G., Winge, P., Medici, S., & Hood, W. M. (2023). Carbohydrate
nutrition associated with health of overwintering honey bees. Journal of
Insect Science, 23(6), 16. https://doi.org/10.1093/jisesa/iead079
Rao, P. V., Krishnan, K. T., Salleh, N., & Gan, S. H. (2016). Biological
and therapeutic effects of honey produced by honey bees and stingless bees: A
comparative review. Brazilian Journal of Pharmacognosy, 26(5), 657–664. https://doi.org/10.1016/j.bjp.2016.03.011
Honey bees (Apis mellifera) rely on a
complex diet to maintain colony health, productivity, and resilience. Among the
most vital components of their nutrition are proteins and carbohydrates. These
macronutrients not only fuel daily activity but also influence brood
development, immunity, longevity, and foraging efficiency. Understanding their
roles and dietary sources is essential for both traditional and modern
beekeeping, especially as environmental changes and habitat loss continue to
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