Bees

Adult bees

There are three adult bees present in bee colonies, at least during the active season. There is usually a single queen and thousands of workers. Differentiation of these two female forms is termed the castecaste:
reproductive division of labor within female bee adults; a single reproductive female queen (via pheromone secretions) maintains more or less sterile female workers
system. There are also male bees—drones—in colonies, except during periods of poor or no incoming resources, such as the overwintering period, when most colonies do not have adult drones.

Worker bees

The hive is populated by female worker bees. Worker bees develop in 21 days. During the adult life span of the worker they perform different tssks: initially hive duties and then collecting resources as a field bee. The worker will pass through a sequence of duties (also called division of labor or temporal castecaste:
reproductive division of labor within female bee adults; a single reproductive female queen (via pheromone secretions) maintains more or less sterile female workers
condition) as they age based on ability to learn and accomplish increasingly complex tasks and on gland development in their adult bodies.

The adult workers initially work within the hive, passing through a sequence of duties based on their age and gland development. Major activities include cleaning, feeding the brood (larvae), storing resources, producing beeswax and building the comb, guarding, and foraging. There are additional specialized tasks involving smaller numbers of worker adults.

Worker adult; photo by Gainenews
Darkly colored worker adult; photo by The BeeMD photo collection
 

Sequence of worker hive duties

Adult bee emerging from cell; photo by The BeeMD photo collection

Newly emerged adults stay close to the brood area, where they emerge from their cappedcapping:
the covering that bees add over comb cells containing fully ripened honey or to cap brood that has reached the pupal stage; bee bread cells are not capped
cells. Their first activity is to clean the cells of the brood area to ready them for the queen to repopulate with an egg. They remove the remains of cappings, polish the cell interior, and fix the cell rim with wax salvaged from cappings. During this time, they consume bee bread, an important behavior critical to complete development of head glandshead glands:
nurse-age bee glands that produce worker jelly or royal jelly. which is fed to developing larvae; these glands are usually mandibular and hypopharyngeal glands
that will be necessary to produce brood food for developing larvae.

Nurse bees are younger workers who tend to the larval cells. They produce food in glands in their heads, which they then put in the bottom of a cell for the larva to feed upon; they do not feed larvae directly. This food is called worker jelly when fed to young (up to three larval days of age) developing workers, or royal jellyroyal jelly:
a protein-rich secretion of the worker's hypopharyngeal glands fed to all young bee larvae. Sometimes termed worker jelly to when it is the food fed to the worker larvae for the first three days of worker larval development; the queen receives this food throughout her larval development
when fed to a developing queen.

Brood cells with younger larvae are supplied with a large amount of jelly, termed “mass provisioning.” Cells with older worker larvae get less food (termed “progressive provisioning”) and lower quality food, which is a mixture of honey and bee bread. Cells are visited up to an estimated 15,000 times during this process. 

Worker adults covering brood, though there’s an inadequate number to keep brood at proper temperature; photo by Rob Snyder
Strong adult population; photo by The BeeMD photo collection
Good worker bee population; arrows indicate bee bread stores; photo by Dewey M. Caron
 

Food processing workers tend to the incoming pollen and nectar.

Comb construction is an activity that is keyed to fully developed wax glands of the worker. Newly established colonies require new comb construction, while mature colonies with frames mainly of drawn comb only need wax to cap brood or cells of fully ripened honey. Thus, the number of workers secreting and molding wax at any one time is dependent on the hive’s needs.

Guarding is worker activity at colony entrances. Guard bees stand high on middle and hind legs with antennaeantenna:
paired, slender, and jointed segmented appendages on the bee head; primary taste, touch and smell receptors. Antennation refers to how bees interact using their antennae to communicate various messages such as food exchange and distribution of queen pheromones.
outstretched at hive openings, just inside the entrance of a hive. They monitor the activity of returning foragers and defend against robbing bees, wasps, or other animal intruders by smelling and watching incoming traffic. If smell or behavior is not characteristic to the colony, an intruder may be challenged. They attempt to grasp and/or block entry of potential intruders.

Despite guards, there is considerable drifting of bees from one colony to another within and even between apiaries; why some bees are allowed to pass while others are challenged is not known. One colony may be invaded by wasp scavengers such as hornets and yellowjackets, or by robbers from another colony. Guards become overwhelmed despite much fighting between intruders and hive occupants. Weaker colonies can be robbed of their food stores and have most of their adult population killed by robbing behavior.

As workers age, they begin to perform orientation flights. Initially, they simply become airborne in front of their colony entrance. Eventually they fan out and learn landmarks to memorize their hive location. During this learning phase, many bees will drift from one colony into another; drifting bees are usually accepted by guard bees. Individuals are thought to take several exploratory flights to learn the site of their home.

Orientation flights are distinctive. They usually occur in the early afternoon hours. Bees leave the landing board and fly in a bouncing, roughly figure 8 motion before their hive. They quickly then move further away, sometimes circling the hive itself. After a period of one to several days, they then become foragers.

Field duties

Foraging represents the last third of a worker bee’s life. The oldest worker bees leave the hive to forage during the last two to three weeks of their normal six-week summer life span. During the winter, their life span is extended to as much as six months when foraging and rearing of brood inside the hive is reduced.

Worker forager with tattered wings; photo by Kathy Keatley Garvey

Foragers support the hive by securing the four resources colonies require: nectar, to ripen into honey; pollen, to process into bee bread; water, to dilute honey and cool or control hive humidity; and propolis, the bee glue that contains antimicrobial properties and helps keep the colony healthy.

Heavily laden foragers may land in the grass, on the ground, on the sides of their hives, or on other structures on their return to the hive. This may also occur when entrance foraging traffic is especially heavy or the entrance is temporarily blocked during beekeeper colony inspection. Foraging needs to be distinguished from heavy entrance activity of orientation flights, from drone flights in afternoons for mating, or when bees from other hives might be robbing.

Size of the primary entrance at the bottom board may interfere with foraging when it is reduced to keep mice out and help bees during the colder winter season. Bees may partially close entrances with propolis, especially those above the bottom landing board entrance. Foragers can be seen coming and going from reduced entrances and at alternative entrances (provided by beekeepers) or from holes or cracks in the hive equipment.

If there is vegetation growth in front of the entrance, foragers may have difficulty when returning to their hive. Beekeepers often provide a landing board before the colony entrance to aid foragers. Use of pollen traps, entrance reducers, robbing screens, or other devices may interfere with forager entry and exiting.

Foragers learn where their hive is via orientation flights as they begin the third week of their adult life. They may discover forage sites on their own as scouts or they may follow directions to potential resource sites provided by dancing bees. Bee dancing is repeated movements of bees following their return form a foraging trip. Dancing conveys specific information about location relative to the sun’s position, distance via energy expenditure, and provides taste and smell clues to assist recruitsrecruits:
older forager-aged bees (generally at least three weeks old) responding to dancing bees seeking to share information about nectar, pollen, water, or bee resin resources
in making foraging decisions. Most foragers work only one or a very limited number of flowering plants. Experienced foragers usually end up with tattered and torn wings.

Some specialized worker duties

Grooming behavior: Worker bees form a queen retinue, continuously grooming her. With this behavior they pick up the queen’s pheromones. Workers also autogroom, using their leg hairs to comb pollen from their body hairs. Workers have a special structure on the front pair of legs to clean the antennaeantenna:
paired, slender, and jointed segmented appendages on the bee head; primary taste, touch and smell receptors. Antennation refers to how bees interact using their antennae to communicate various messages such as food exchange and distribution of queen pheromones.
. Workers also groom other worker bees (allogrooming). Allogrooming helps remove varroa mites. Workers also groom drones.

Undertaker bees: Bees die within the hive continually. Undertaker bees find these dead bees, some of which fall to the bottom board, and remove their bodies from the hive and bottom board. Several bees might be involved in removing a single dead bee body.

Heater bees: During cluster behavior, some bees crawl into brood cells and elevate their body temperature by moving their wing muscles without moving their wings. Heat radiates from their body, which serves to help adjacent developing pupae develop normally and adds to the heat retained by the cluster.

Resources

Delaplane KS. 2019. Biology of Individual Honey Bees. Bee Health. Accessed 2024. https://bee-health.extension.org/biology-of-individual-honey-bees/?msclkid=c3cb798ccfcc11ecb5773b5bfd6f1e2d

Caron DM and Connor LJ. 2022. Honey Bee Biology and Beekeeping. Wicwas Press, Kalamazoo, MI, US.