Perspectives on hygienic behavior in Apis mellifera and other social insects
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Apidologie * The Author(s), 2020 DOI: 10.1007/s13592-020-00784-z
Perspectives on hygienic behavior in Apis mellifera and other social insects Marla SPIVAK1 , Robert G. DANKA2 1 Department of Entomology, University of Minnesota, 1980 Folwell Ave., St. Paul, MN 55018, USA Honey Bee Breeding, Genetics, and Physiology Laboratory, USDA-ARS, 1157 Ben Hur Rd., Baton Rouge, LA 70820, USA
2
Received 26 February 2020 – Revised 14 May 2020 – Accepted 5 June 2020
Abstract – Hygienic behavior in honey bees, Apis mellifera , has been studied for over 80 years with the aim of understanding mechanisms of pathogen and parasite resistance and colony health. This review emphasizes the underlying behavioral mechanisms of hygienic behavior in honey bees and when known, in other social insects. We explore the relationship between honey bee hygienic behavior toward diseased brood and Varroa- parasitized brood (Varroa -sensitive hygiene, VSH); the timing of hygienic removal of diseased, Varroa -infested, and virus-infected brood relative to risk of transmission that can affect colony fitness; and the methods, utility, and odorants associated with different assays used to select colonies for resistance to diseases and Varroa . We also provide avenues for future research that would benefit honey bee health and survivorship. social immunity / Varroa -sensitive hygiene / chemical odorants / ants / termites
1. INTRODUCTION Hygienic behavior is an important form of social immunity (Cremer et al. 2007) for a number of social insect species. The term hygienic behavior was coined by Rothenbuhler (1964) to describe the process of detection and elimination of diseased brood by adult honey bees (Apis mellifera ). The term “Varroa -sensitive hygiene” (VSH) was coined more recently (Harris 2007) to describe the detection and removal of brood infested with the parasitic mite Varroa destructor by honey bees (Harbo and Harris 2005). The behavioral sequence of uncapping and removing the brood, as first described (Rothenbuhler 1964), is the same whether the brood is diseased, mite-infested, or dead, but this motor pattern may be triggered by the detection of Corresponding author: M. Spivak, [email protected] Manuscript editor: James Nieh
different odorants associated with the health status of the brood. In honey bee colonies, elimination of brood consists of adult bees removing and/or cannibalizing the abnormal brood from individual cells, either intact or in pieces, and discarding remains outside the hive; in Reticulitermes termites, it consists of cannibalization (Davis et al. 2018) and in Lasius ants of destructive disinfection by dismembering the infected pupa and then disinfecting with venom (Pull et al. 2018). Hygienic behavior helps maintain the health of densely populated insect societies by limiting horizontal transmission of pathogens and population growth of parasites. Workers that destructively eliminate already infected or infested individuals protect the colony, or superorganism, in a similar way to immune cells that protect an organism f
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