Elemir Simko, Sarah Wood, and Ivanna V. Kozii In North America, antimicrobials have been used for decades as part of integrated pest management of infectious disease in honey bees. In contrast, in the European Union, it is illegal to treat honey bee colonies with antimicrobials. Since 2017 in the United States, and 2018 in Canada, veterinarians have been federally mandated to oversee prescribing of medically important antimicrobials to beekeepers for treatment of their colonies. Considering that many veterinarians do not receive education in honey bee medicine and disease, this chapter provides guidance for veterinarians who wish to better serve their beekeeper clientele. A basic understanding of honey bee biology is essential to understanding pathogen transmission and disease control in honey bees. Honey bees are social insects. Under normal circumstances, a honey bee colony consists of three castes: a queen, drones, and worker bees. Both queen and workers are diploid females and have 32 chromosomes, while drones are haploid males with 16 chromosomes. The honey bee queen is the only mated, egg‐laying female with a developed reproductive tract in the colony. The queen can lay ~1500 eggs per day that develop into workers, drones or queens (if needed). Besides egg laying, the other major role of the queen is production of multiple pheromones that maintain colony homeostasis and productivity (Bortolotti and Costa, 2014). It is common for beekeepers to replace the queen in their colonies every two years to ensure vigorous egg and pheromone production, which in turn promotes colony health and vitality. Drones are male bees whose main function is to mate with virgin queens, after which the drones die. During the summer, drones often drift from colony to colony, providing an opportunity for pathogen transmission among colonies (Table 40.1). During the fall, surviving drones are expelled from the colony in preparation for winter. Worker honey bees are sterile females that have hypoplastic ovaries. Worker honey bees are responsible for all activities in the colony except reproduction. The honey bee workers are divided into two major types based on seasonality and tasks: summer and winter workers. Summer workers live approximately six weeks. The first half of the life of an adult summer worker (~first three weeks) will be dedicated to housekeeping tasks in the hive, including cleaning brood cells; nursing brood; attending to and feeding the queen; building wax comb and sealing brood and honey; processing nectar into honey; processing pollen into bee bread; removing dead brood and adults from the hive; and ventilating and defending the colony. Many of these behaviors facilitate horizontal transmission of pathogens within a colony (Table 40.1). The second half of an adult summer worker’s life (~lasts three weeks) is dedicated to field work which consists of collecting nectar, pollen, water, and propolis. The most intense foraging occurs within two miles, but the forager bees can fly twice as far if nectar or pollen are not available closer. Drifting of honey bee workers from hive to hive is not as common as in drones but it does occur. Table 40.1 Behaviors important in horizontal spread of pathogens of honey bees. Winter workers emerge at the end of summer and the beginning of fall. Their life span is 150 days on average (Currie et al., 2015). The main role of winter worker bees is to ensure that the queen survives until the spring season begins and a new population of bees is generated. The winter bees usually do not defecate within the hive; instead, they accumulate waste within the rectum which can distend and occupy a substantial portion of the abdominal cavity. On sunny winter days when the temperature is above 0° C, bees will fly out of the hive for a short period to empty the waste from their rectum (cleansing flights). There are seasonal variations in the total number of worker bees per colony. To maximize honey production, the highest number of worker bees in a colony (50,000–70,000 bees) should coincide with the major blooming season of nectar‐producing plants. In contrast, there are only 15,000–30,000 bees in a colony during the late winter. Developmental stages of brood are the same for all three castes, even though the total development time is different for each caste. From fertilized eggs, diploid larvae emerge three days after oviposition and will develop into workers or queens depending on the larval food they are provided by nursing bees. Nonfertilized eggs will develop into haploid drones. After hatching, larvae are attended and nursed by many young nursing bees visiting each larva many times per day. After completion of the larval stage (5–6 days), the comb cells containing larvae are sealed with wax (capped) and prepupal and pupal development occurs for 8–15 days, depending on caste. Eclosion (emergence) of the imago (adult) stage occurs at day 16, 21, and 24 post oviposition for queen, worker, and drone, respectively. Larvae and pupae that die during the development process are removed and their brood cells are cleaned by housekeeping worker bees. In addition, worker bees have the ability to detect diseased pupae and remove them; this is called “hygienic behavior,” which interferes with disease progression and improves colony resistance to disease. Hygienic behavior is a part of social immunity and contributes significantly to the overall health of a colony. This is an inherited trait and there are various queen‐breeding programs that selectively enhance this behavior. A honey bee hive contains thousands of bees with biological behavior that requires close interaction (e.g., food exchange or trophallaxis), direct contact (e.g., pheromone spread), and housekeeping duties (e.g., removal of dead brood and adult bees). These behaviors facilitate horizontal transmission of pathogens between individual bees (Table 40.1). Once a disease is established within a colony, it can spread from colony to colony by natural or anthropogenic means. Robbing behavior is the most important natural mode of transmission of honey bee diseases between colonies. Robbing occurs when forager bees find an unprotected source of honey that is collected and brought back to their hive. Unprotected stores of honey are often available in colonies that are dead or weak due to disease; accordingly, pathogens from diseased colonies are transmitted to healthy and strong colonies by their strong foraging population (robber bees). Anthropogenic transmission of disease occurs when beekeepers spread pathogens through the use or sale of contaminated bees, hive products, or equipment. In North America, antimicrobials are used as part of an integrated pest management program for control of disease in honey bees caused by bacterial and microsporidian pathogens. These diseases include American foulbrood, European foulbrood, and Nosema disease. American foulbrood (AFB) is a devastating, contagious brood disease that develops rapidly, kills the colony and spreads to other colonies by robbing, drifting bees, and anthropogenic modes. It is caused by Paenibacillus larvae, a Gram‐positive, spore‐forming, rod‐shaped bacterium. Spores survive in contaminated equipment for decades. Approximately 2.5 billion spores are produced in each infected larva (Pettis, 2015). Larvae (up to two days old) ingest spores which germinate and proliferate in the midgut and subsequently spread throughout the body, causing fatal septicemia (Pernal, 2013; Pettis, 2015). Affected brood frames have a spotty brood pattern (shotgun brood) with perforated and/or sunken capping of brood cells (Figure 40.1A); dead, dull white to brown pupae that are positive for the “ropiness” test (Figure 40.1B); and desiccated, dead pupae which form dark brown, brittle scales firmly adhered to the “ventral lateral” wall of the brood cell (scale cannot be removed without destroying the cell wall; Figure 40.1C,D). The large number of vegetative P. larvae bacteria within macerated dead pupae will generate a typical glue‐like consistency that can be detected by the “ropiness” test. To perform the test, a dead pupa is macerated with a matchstick within a cell and then slowly withdrawn. If the macerated tissue can be drawn out and stretched more than 2 cm, it is indicative of AFB infection (Pernal, 2013; Pettis, 2015). In advanced stages of disease there may be a strong decaying odor when the colony is opened, hence the name of the disease, “foulbrood” (Pernal, 2013; Pettis, 2015). Gross pathology of the brood frame, especially a positive “ropiness” test (Figure 40.1B) and the presence of scales (Figure 40.1C,D), is highly characteristic, or could be considered even pathognomonic, for AFB. Nevertheless, submission of samples of affected brood (including scales, if present) is recommended for confirmation of diagnosis by bacterial culture and/or PCR. Importantly, veterinarians suspecting AFB should contact their state/provincial apiarist and/or veterinarian, as management and control of AFB are subject to state/provincial regulations and AFB is a notifiable disease in some jurisdictions (Pernal, 2013; Pettis, 2015).
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Antimicrobial Therapy in Honey Bees
Introduction
Honey Bee Biology
Group/caste
Behavior
Drone, worker
Drifting among colonies
Worker
Removing dead brood and adults
Cleaning brood cells
Nursing and feeding brood
Attending to and feeding the queen and drones
Processing pollen into bee bread
Processing nectar into honey
Foraging
Exchanging food among workers (trophallaxis)
Robbing
Drone, worker, queen
Close contact and interaction
Beekeeper
Use or sale of contaminated bees, hive products, or equipment
Transmission of Honey Bee Disease
Honey Bee Pathogens and Diagnosis
American Foulbrood

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