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Laying worker bee
Laying worker bee
Laying worker bee
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Laying worker bee honeycomb. See broad pattern and drone brood in worker cells (caps protruding). This honeycomb is taken from the dying family without the queen.

A laying worker bee is a worker bee that lays unfertilized eggs, usually in the absence of a queen bee. Only drones develop from the eggs of laying worker bees (with some exceptions, see thelytoky). A beehive cannot survive with only a laying worker bee.[1]

Prevalence

[edit]

Even in a normal hive, about 1% of workers have ovaries developed enough to lay eggs. However the usual number of the laid eggs is very small. Only eight eggs (seven moderately and one fully developed) were found after examining of 10,634 worker bees[2] (strong colony contains about 100,000). Workers eventually lay significant numbers of eggs only in queenless colonies.

Development

[edit]

Laying workers develop in the absence of open brood as produced by a healthy adult queen. Normally, pheromones from the brood – known as brood recognition pheromones – prevent development of the workers' ovaries. Laying workers can develop after the colony's queen has been lost to swarming,[3] or in the presence of a failing queen which has yet to be superseded. The process of developing a laying worker usually takes weeks after the loss of the original queen. In adult laying workers there is an anatomic (and physiological) trade-off between the sizes of their more developed ovaries and their less developed food glands.[4][5]

Identification

[edit]

All methods of identifying a laying worker bee involve inspection, in which the beekeeper examines the brood pattern and type to identify if a healthy queen is present, or a potential laying worker. The beekeeper looks for diagnostic signs, including:[citation needed]

  • Brood pattern
    Laying workers lay eggs that lack the queen's egg recognition pheromone, meaning that other workers may remove the eggs. This results in a spotty brood pattern, in which empty cells are scattered heavily through capped brood.
  • Number of eggs per cell
    The beekeeper looks at the honeycomb cells to see how many eggs are laid in each one. Queen bees will usually lay only a single egg to a cell, but laying workers will lay multiple eggs per cell. Multiple eggs per cell are not an absolute sign of a laying worker because when a newly mated queen begins laying, she may lay more than one egg per cell.
  • Egg position
    Egg position in the cell is a good indicator of a laying worker. A queen bee's abdomen is noticeably longer than a worker, allowing a queen to lay an egg at the bottom of the cell. A queen bee will usually lay an egg centered in the cell. Workers cannot reach the bottom of normal depth cells, and will lay eggs on the sides of the cell or off center.
  • Drone brood in worker cells
    Another good indicator is drone brood in worker sized cells. Drones are raised in larger cells than workers. Drone cells are recognizable by their larger size; and when capped, drone cells are capped with blunt pointed cappings. Drones in worker cells are a sure sign of a failing queen or laying worker.

References

[edit]
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Laying worker bee

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A laying worker bee is a female worker honey bee (Apis mellifera) whose ovaries become activated to lay unfertilized eggs, resulting in the development of haploid male drones rather than diploid female workers or queens. These eggs are typically deposited in worker cells, leading to undersized but functional drones that emerge after about 24 days. This reproductive behavior represents an adaptive response in queenless colonies, where the absence of queen mandibular pheromone (QMP) fails to suppress worker ovarian development. Laying workers arise primarily when a colony becomes queenless for an extended period, often due to queen loss, failure, or supersedure issues, allowing the buildup of worker-laid eggs over time. In such scenarios, not all workers activate ; only a subset—typically younger adults—develop functional ovaries, influenced by factors like colony size and the presence of open brood that normally inhibit this process. The phenomenon can also occur in colonies with diseased or aging queens, where signals weaken, mimicking queenlessness. Biologically, laying workers exhibit physiological changes, including upregulation of vitellogenin genes for production and enhanced stress resistance, which may prolong their lifespan compared to non-reproductive workers. However, their has detrimental colony-level effects: the production of drones consumes resources without contributing to or tasks, leading to reduced worker populations, foraging efficiency, and overall hive productivity. In unmanaged cases, this can culminate in colony decline or , though —where other workers remove and consume laid eggs—helps mitigate the issue in queenright hives. In beekeeping, early detection via multiple eggs per cell or spotty drone brood patterns is crucial for intervention, such as requeening or adding brood to restore pheromone balance.

Definition and Biology

Physiological Characteristics

In normal worker honey bees (Apis mellifera), the ovaries are underdeveloped and contain a fixed number of 2–12 ovarioles per , which is substantially fewer than the 150–200 ovarioles per found in . These ovarioles remain inactive under typical queenright conditions due to inhibitory pheromones, preventing egg production. In laying workers, which develop in queenless colonies, the existing ovarioles activate and mature, enabling the production of unfertilized eggs, though the total ovariole count does not increase post-emergence. This activation results in ovaries that are enlarged compared to sterile workers but remain smaller and less productive than those of . The physiological shift to laying in workers involves distinct hormonal profiles, including low titers of (JH) in the , similar to those in and nurse bees, rather than the elevated JH levels associated with foraging. This low JH state correlates with ovary activation and supports reproductive behavior in laying workers. Additionally, vitellogenin levels, a key yolk precursor protein, are elevated in the of laying workers, facilitating and distinguishing them from foraging workers, which exhibit suppressed vitellogenin synthesis. Queen pheromones normally suppress these changes by inhibiting JH modulation and vitellogenin expression, but their absence allows reproductive physiology to dominate. Laying workers exhibit physiological trade-offs due to energy reallocation toward , including a delayed transition to and reduced engagement in collection compared to sterile workers. This shift prioritizes ovarian development over tasks like or resource gathering, leading to shorter lifespans and diminished colony-level efficiency. Hypopharyngeal glands in laying workers often remain active or more developed than in typical foragers, allowing dual roles in brood food production and egg-laying, though overall gland function may decline with age as reproductive demands intensify. Laying workers produce only unfertilized eggs, which develop into haploid males (drones) via , contributing minimally to colony reproduction since no females result. An exception occurs in thelytokous populations, such as the (Apis mellifera capensis), where laying workers can produce diploid female offspring through automixis, potentially leading to social .

Developmental Process

In queenless honey bee (Apis mellifera) colonies, the developmental process of laying workers begins with the cessation of queen mandibular pheromone (QMP) and brood pheromone suppression, which normally inhibits worker activation. This inhibition release triggers initial physiological changes within days of queen loss, including the upregulation of vitellogenin synthesis for yolk deposition in oocytes. Early stages involve partial enlargement from undeveloped states (<0.8 mm) to intermediate development (0.8–1.2 mm), typically detectable 5–10 days post-queen removal, as workers transition from nursing roles to reproductive activation. Oocyte maturation follows, progressing to fully developed ovaries (>1.2 mm) capable of production, with peak development occurring around 28–31 days after queen loss. Mass egg-laying capability emerges abruptly between 22–37 days (mean 28.9 days), when workers deposit unfertilized haploid s primarily in drone cells. This timeline aligns with the exhaustion of existing brood, amplifying pheromone deficits and accelerating the process. Genetic factors, such as variations, influence individual propensity, while environmental cues like and modulate progression speed. Younger workers, particularly 4–8-day-old nurse bees, are most prone to rapid development due to their physiological readiness for vitellogenin production and trophallactic interactions that provide essential proteins. In contrast, older foragers exhibit minimal activation. Colony size also plays a key role: smaller groups (25–125 workers) promote faster and higher rates of development compared to larger populations (e.g., 600 workers), likely due to reduced for resources and intensified dilution. Up to 5–24% of workers may achieve full fertility in such conditions. Once ovaries fully develop and egg-laying commences, the transformation is largely irreversible; even upon queen reintroduction, resorption or may reduce activity, but activated workers rarely revert to complete sterility, perpetuating low-level in the .

Causes and Occurrence

Triggers in Colonies

In colonies, the development of laying workers is primarily triggered by the absence of key inhibitory s, particularly queen mandibular (QMP) and brood s, including the brood ester (a blend of 10 esters) and the volatile primer E-β-ocimene. QMP, produced by the queen's mandibular glands, normally suppresses ovarian development in workers by modulating hormonal pathways, including reduced (JH) titers and induction of in ovarian cells, thereby preventing reproduction. Similarly, brood s emitted by open larval brood inhibit worker activation through comparable hormonal mechanisms, reinforcing reproductive restraint in the presence of a functional queen and brood. When these s are absent, workers experience unrestrained ovarian development, leading to egg-laying behavior as a facultative response to perceived reproductive failure. Common scenarios initiating this process include sudden queen death, which abruptly halts QMP production; swarming events, where the departing swarm leaves the parent colony temporarily queenless; and failed queen rearing, such as when emergency queen cells fail to produce a viable mated queen. These events are often exacerbated in small or stressed colonies, where distribution may be less effective. The inhibitory effects of QMP and brood pheromone operate through integrated neural and hormonal pathways in worker bees. QMP influences brain levels, which in turn regulate behavioral and reproductive states, while both pheromones downregulate vitellogenin expression and JH synthesis in the corpora allata, directly suppressing . Historical observations, such as those documented by Seeley in studies of pheromone dynamics, highlight how queenless hives rapidly shift to worker when these suppressive signals diminish, underscoring the pheromones' role in maintaining colony-level reproductive control.

Prevalence and Frequency

In queenright colonies of the Apis mellifera, fewer than 0.01% of workers exhibit fully activated ovaries capable of egg production, though actual instances of worker egg-laying remain exceedingly rare due to policing behaviors by other workers. For instance, dissections of 10,634 workers from 21 colonies revealed only seven with moderately developed eggs (about half the size of mature ones) and none with fully mature eggs ready for laying. This low frequency underscores the strong suppression of worker under normal conditions, where queen pheromones inhibit ovarian development in the vast majority of workers. In contrast, queenless colonies see a marked increase in laying workers, with a significant proportion of workers, up to 80–90% in some studies, developing active ovaries after several weeks of queen loss. This proportion can rise even higher in small, isolated, or resource-limited hives, where the absence of brood pheromones accelerates ovarian activation among younger workers. The progression typically starts with a small of pseudoqueens (reproductively primed workers) that lay eggs haphazardly, often multiple per cell, leading to a feedback loop that recruits more workers into laying roles. Prevalence varies across subspecies and strains. In Africanized honey bees (A. mellifera scutellata hybrids), worker reproduction contributes more substantially to drone production, with laying workers playing a larger role in colony fitness due to higher rates of ovarian activation and reduced policing efficiency compared to European strains. Notably, in the (A. mellifera capensis), laying workers exhibit —a form of —allowing their unfertilized eggs to develop into diploid females rather than solely haploid drones, which can sustain or parasitize colonies in rare cases. This trait, controlled by a single recessive locus, occurs infrequently outside capensis but highlights genetic influences on worker laying frequency. While foundational studies from the – provide these benchmarks, quantitative data on laying worker prevalence remain sparse in post-2012 literature, with limited exploration of modern stressors like mite infestations potentially exacerbating queenlessness and thus worker laying in stressed colonies. Recent research (2020–2025) has focused more on strategies, such as curing laying workers in new divisions and their behavior in queen monitoring cages, but has not substantially updated prevalence estimates.

Impacts and Consequences

Effects on Hive Dynamics

The presence of laying workers induces significant behavioral shifts within the colony, disrupting the established division of labor. These workers often produce sounds similar to those emitted by , with fundamental frequencies around 350 Hz, which can elicit confused responses from other bees and lead to attacks on the piping individuals. Additionally, laying workers produce secretions from their Dufour's gland that mimic components of the , attracting a of attendant workers who groom and feed them, thereby diverting colony resources toward non-reproductive individuals. This mimicry contributes to a broader breakdown in task allocation, where affected workers, including potential laying workers, reduce engagement in brood or for and , reverting to more ancestral, less specialized behaviors that diminish overall hive efficiency. These disruptions escalate into social conflicts, as the mounts a defense against selfish reproduction through heightened . Other workers aggressively remove worker-laid eggs, achieving removal rates approaching 100% in queenright colonies of certain strains like Apis mellifera carnica, to enforce sterility and favor queen-produced offspring. In queenless conditions, policing persists but may be less effective. Such conflicts highlight the tension between and interests, intensifying intra-colony strife. The cumulative effects accelerate decline, particularly in queenless where laying workers predominate. Resources that would normally support brood rearing and maintenance are redirected toward the care of laying workers and their infertile offspring, leading to neglected hive , depleted stores, and heightened vulnerability to during lulls. Queenless with laying workers also exhibit faster progression to disease susceptibility, as weakened reduces the influx of and . These dynamics often result in irreversible hive collapse within weeks if no new queen is introduced. From an evolutionary standpoint, the emergence of laying workers exemplifies a in eusociality, where individual workers pursue personal reproduction at the colony's expense, undermining the inclusive fitness benefits that stabilize cooperative societies. This conflict arises because workers are more closely related to queen-laid sisters (r=0.75) than to their own sons (r=0.5), favoring policing to promote kin-selected as outlined in Hamilton's theory.

Reproductive Outcomes

Laying workers in honey bee colonies (Apis mellifera) produce unfertilized, haploid eggs that develop exclusively into male drones, as these workers lack the ability to mate and fertilize their eggs. This reproductive limitation ensures no female offspring—workers or queens—are generated from standard laying worker eggs, severely restricting colony propagation. An exception occurs in the Cape honey bee subspecies (A. m. capensis), where laying workers exhibit thelytokous parthenogenesis, allowing unfertilized eggs to develop into diploid females that can become workers or queens. This rare trait, controlled by a single major gene, enables pseudocloning but is not observed in most honey bee populations. The brood patterns resulting from laying workers are distinctive and disruptive. Multiple eggs, often numbering 2 to 10 or more, are commonly laid in a single cell due to the uncoordinated efforts of several laying workers, leading to frequent supersedure where stronger larvae consume weaker ones or to outright of excess eggs by nurse bees. Additionally, drone brood appears in worker-sized cells, creating a spotty, irregular across the that contrasts with the uniform worker brood produced by a queen. Genetically, the offspring of laying workers contribute to reduced diversity within the colony's reproductive output. All drones produced are direct sons of the laying workers, who themselves originate from the colony's original queen, resulting in a narrow genetic pool that promotes potential if these drones were to mate with related from nearby colonies. This limitation exacerbates the colony's vulnerability, as no new female bees are produced to sustain workforce replacement. In the long term, colonies dominated by laying workers inevitably fail without intervention to introduce a queen. The absence of worker brood means the existing population dwindles as foragers age and die, with the typically collapsing within 2 to 4 months due to the unsustainable production of only drones.

Detection and Management

Identification Methods

Beekeepers can identify laying worker bees primarily through of the brood frames during hive examinations. A key indicator is the presence of spotty or irregular brood patterns, where capped brood appears scattered with numerous empty cells interspersed, unlike the compact, pattern produced by a laying queen. This irregularity arises because laying workers deposit eggs haphazardly across the . Another prominent visual cue is multiple eggs per cell, often two to several, which are typically laid off-center, on the side walls, or even on the top of the cell, contrasting with the single, upright placed at the bottom by a queen. Examination of the brood further confirms the presence of laying workers. Worker-laid eggs develop exclusively into drones, resulting in drone brood—characterized by larger, domed cappings—in worker-sized cells, often scattered throughout the frame rather than in dedicated drone areas. Additionally, the absence of queen cups or other signs of an active laying queen, such as a solid patch of worker brood, supports this diagnosis, as laying workers do not stimulate the colony to rear queens. To differentiate from a drone-laying queen, note that the latter produces a more organized pattern with fewer multiple eggs. Behavioral observations provide supplementary evidence. Laying worker colonies often exhibit increased aggressiveness during inspections, with bees displaying heightened defensive responses due to the lack of queen pheromones suppressing worker ovarian development. Laying workers themselves may appear with slightly rounded or developed abdomens from ovarian activity, though this is subtler than a queen's elongated form and requires close scrutiny. Drone brood traps, consisting of frames with foundation designed for worker cells, can be introduced to the hive; the subsequent appearance of drone brood in these cells within a few weeks indicates laying worker activity. Practical tools and techniques enhance detection accuracy. During inspections, gently shaking a frame can dislodge bees to reveal eggs or larvae more clearly, allowing beekeepers to spot multiple eggs or irregular placements without obstruction. Timing is crucial: laying workers typically emerge 2-3 weeks after queen loss, when open brood pheromones diminish, so routine checks around this period post-suspected queenlessness are recommended. These methods, when combined, enable reliable identification while minimizing hive disturbance.

Prevention and Control Strategies

Prevention of laying worker development primarily involves vigilant monitoring and timely intervention to maintain queenright status in the . Beekeepers are advised to conduct regular inspections, ideally every 7 to 10 days during active seasons, to verify the presence of a healthy laying queen and detect queenlessness early. If queen loss is suspected, introducing a new mated queen or a frame containing eggs and young larvae (less than three days old) within two to three weeks of queenlessness can suppress worker ovarian development through the reintroduction of queen mandibular pheromone (QMP) and brood pheromone, preventing the onset of laying workers. Annual or biannual queen replacement, particularly in commercial operations, further reduces the risk by ensuring consistent pheromone production and stability. Additional preventive measures focus on minimizing colony stress that could lead to queen failure or loss. (IPM) practices, such as monitoring and treating for mites with approved miticides during peak infestation periods, help maintain overall hive health and reduce stressors that exacerbate queenlessness. Avoiding overcrowding by timely supering or splitting strong colonies also prevents swarming and subsequent queen issues, while providing adequate nutrition through supplemental feeding during dearth periods supports queen productivity. Once laying workers are established, control strategies aim to suppress their reproduction and restore queenright conditions, though success diminishes if the condition persists beyond a few weeks. A common method is the addition of open brood frames from a donor queenright , introduced every five to seven days for two to three weeks, to expose the hive to brood pheromone (E-β-ocimene), which inhibits worker ovarian activation and reduces laying worker activity. Following this suppression, requeening can be attempted using a strong nucleus via the newspaper combine method, where the is placed atop the laying worker hive separated by newspaper, allowing gradual pheromone integration; early intervention yields success rates of 70-90% in accepting the new queen. Alternatively, queen pheromone lures can be used adjunctively to mimic QMP and aid acceptance during requeening. The shook swarm technique offers another elimination approach by shaking the laying worker bees into a new hive location on a sunny day after has begun, often at the apiary edge, where poorly oriented laying workers fail to return while foragers reorient; the cleaned equipment is then provided with foundation and brood from a queenright source before requeening. To support requeening, drone brood should be culled by scraping or replacement to eliminate reinforcing cues for worker laying. In cases of advanced decline, combining with a strong queenright using a double-screen board for can integrate the bees while suppressing laying workers, though direct culling of the affected may be necessary if recovery efforts fail. Post-2020 apiculture guidelines emphasize combining these methods with IPM to enhance long-term resilience.

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