Bee Lab

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Climatic limits of tropical African honeybees in the Americas
(International Bee Research Association, 1984) Orley, Taylor R., Jr.; Spivak, Marla
Honey production by Africanized and European honey bees in Costa Rica
(Springer, 1989) Spivak, M.; Batra, S.; Segreda, F.; Castro, A.L.; Ramirez, W.
Seventeen colonies of bees were used in a test of honey production in a coffee plantation in Costa Rica. The identities of the colonies were not known at the beginning of the experiment. Behavioral identifications were made in the field as to whether a colony was ’strongly Africanized’, ’strongly European’ or ’intermediate’. The distance spanned by 10 linear worker cells was also measured in the field. Morphometric analyses, conducted independenUy, identified colonies as probably Africanized or probably European. Although the ’intermediate’ category tended to produce more honey, there was no significant difference in honey production between bee types, whether sorted by behavior or morphometrics. There was no significant correlation between initial colony weight or brood area and total amount of honey produced. Appropriate uses of behavioral and morphometric identifications are discussed.
Do measurements of worker cell size reliably distinguish Africanized from European honey bees (Apis mellifera L.)?
(American Bee Journal, 1992) Spivak, M.; Erickson, E. Jr.
Two experiments were conducted to determine whether the size of the cells in which bees develop affects the size of the cells they subsequently construct. The results indicate that when the size of bees has been modified through the use of foundation with larger or smaller cell bases, bees will construct natural cells of a size consistent with their genetic origin. When 22 Africanized colonies were hived on commercial European foundation, they subsequently constructed · natural cells which were not significantly different in size from those of 66 Africanized colonies hived on all natural comb. Also, eight European colonies reared on noncommercial foundation with small cell bases subsequently constructed natural cells which were significantly larger than the cells from which they emerged. Therefore, averaging the width of 10 linear cells in three diagonal rows on naturally built comb is a relatively reliable and accurate method to distinguish between Africanized and European bees in the field. However, colonies with intermediate cell sizes, which may include some feral European colonies and "hybrids" between Africanized and European colonies, may not be evaluated with certainty.
Facultative expression of hygienic behaviour of honey bees in relation to disease resistance
(Taylor and Francis, 1993) Spivak, Marla; Gilliam, Martha
Four experiments were conducted to examine factors that influence the expression of hygienic and non-hygienic behaviour in honey bees, Apis mel/itera, and to examine the correlation between this behaviour and resistance to chalkbrood, Ascosphaera apis. Colonies were headed by instrumentally inseminated queens selected on the basis of uncapping and removal behaviour expressed by their progeny. In the first experiment, colony strength was altered by transferring hygienic and nonhygienic colonies from 1O-frame field hives to 2-frame observation hives. This treatment significantly reduced the hygienic response of the hygienic bees but did not affect the response of the non-hygienic bees. In the second experiment, hygienic and non-hygienic bees displayed different responses to freeze-killed and live brood which had been partially or entirely uncapped. Both lines of bees recapped both partially and entirely uncapped live brood, but non-hygienic bees also recapped partially uncapped freeze-killed brood, suggesting that non-hygienic bees either could not detect dead or diseased brood or avoided it by sealing it within a comb cell. The third experiment tested whether the degree of hygienic behaviour could be increased by adding hygienic bees to non-hygienic colonies. Adding 20-30% young hygienic bees to nonhygienic colonies did not increase the degree of hygienic behaviour, but adding young nonhygienic bees to hygienic colonies suppressed the behaviour. The results suggest that although hygienic behavior is genetically determined, its expression depends on colony strength and composition of workers within the colony. In the fourth experiment, the hygienic and non-hygienic colonies were fed with pollen patties containing A. apis spores. The weak correspondence that was observed between removal behaviour and physiological resistance to chalkbrood suggested that few colonies are both highly hygienic and physiologically resistant to chalkbrood. Selection against uncapping and removing diseased brood might occur if this behaviour also promotes the spread of disease through the colony. This possibility is discussed in relation to avoidance behaviour of other social insects toward pathogens.
The influence of temperature on cuticular color of honeybee (Apis mellifera L) queens
(Springer, 1993) DeGrandi-Hoffman, G.; Spivak, M.; Martin, J.
Temperatures were monitored around emergency queen cells in queenless honeybee hives to determine the effect of temperature on queen color. Concurrently, sister queens of those reared in the queenless colony were placed in incubators set at 31.1, 32.8, or 34.4 °C for the postcapping interval. Queens from the 34.4 °C incubator were significantly lighter in color than those in the 31.1 °C in all but one trial. Queens that developed in the colony were not significantly different in color rank from those that emerged in the 34.4 °C incubators in any trial, and from the 32.8 or 31.1 °C incubators in most trials. The median color ranks of queens emerging in the colony did not differ throughout the year.
Honey bee hygienic behavior and defense against Varroa jacobsoni
(Springer, 1996) Spivak, Marla
Hygienic and non-hygienic colonies from ’Starline’ stock of Apis mellifera were tested for their ability to remove pupae infested with Varroa mites. The hygienic and non-hygienic lines were selected and bred on the basis of their removal response to freeze-killed brood. A Jenter Box® was used to test whether they would remove experimentally infested pupae following methods of Boecking and Drescher (1992). In 1994, the hygienic colonies removed significantly more pupae infested with one mite per cell than the non-hygienic colonies. In 1995, there was no significant difference between the hygienic and non-hygienic colonies when one or two mites were introduced per pupa due to variation in response among hygienic colonies. There was no significant difference between the rate of removal of infested pupae from the Jenter Box and from natural wax comb by the hygienic colonies. The number of mites damaged by grooming ranged from 6.0 to 42.3% among all colonies. The reproductive success of the mites not removed from the cells by the bees was low in both hygienic and non-hygienic colonies.
Performance of hygienic honey bee colonies in a commercial apiary
(1998) Spivak, Marla; Reuter, Gary S.
Colonies with naturally mated queens from a hygienic line of Italian honey bees (Apis mellifera ligustica) were compared to colonies from a commercial line of Italian bees not selected for hygienic behavior. The following characteristics were compared: rate of removal of freeze-killed brood; amount of chalkbrood; incidence of American foulbrood; honey production; and the number of mites, Varroa jacobsoni, on adult bees. The hygienic colonies removed significantly more freeze-killed brood than the commercial colonies, had significantly less chalkbrood, had no American foulbrood, and produced significantly more honey than the commercial colonies. Estimates of the number of Varroa mites on adult bees indicated that the hygienic colonies had fewer mites than the commercial colonies in three of four apiaries. In previous studies on the relation between hygienic behavior and resistance to diseases and mites, the test colonies contained instrumentally inseminated queens. This is the first study to evaluate hygienic stock in large field colonies with naturally mated queens.
Hygienic behaviour of honey bees and its application for control of brood diseases and varroa: Part I. Hygienic behaviour and resistance to American foulbrood
(International Bee Research Association, 1998) Spivak, Marla; Gilliam, Martha
There have been very few studies on hygienic behaviour as a mechanism of resistance to American foulbrood since Park, Woodrow, Rothenbuhler, and Rothenbuhler’s students published their seminal work.The studies outlined in this part of the review form the core of information from which all later studies on hygienic behaviour have been based.
Hygienic behaviour of honey bees and its application for control of brood diseases and varroa: Part II. Studies on hygienic behaviour since the Rothenbuhler era
(International Bee Research Association, 1998) Spivak, Marla; Gilliam, Martha
Part I of this review summarized the initial research on hygienic behaviour of honey bees, Apis mellifera. This early work that concerned hygienic behaviour as a mechanism of resistance to American foulbrood (AFB) has been the foundation for all subsequent research on hygienic behaviour. In Part II, research on hygienic behaviour in relation to other bee diseases and to Varroa jacobsoni and in Apis species and subspecies is reviewed. In addition, techniques to screen bee colonies for the behaviour are detailed, and practical applications of breeding bees for hygienic behaviour are given. A section on neuroethology demonstrates how modern neurobiological techniques are being used to detect the reasons for differences in responses of hygienic and non-hygienic bees to abnormal brood.
Behavioral defenses of honey bees against Varroa jacobsoni Oud.
(1999) Boecking, Otto; Spivak, Marla
Two behaviors of honey bees, hygienic behavior and grooming, are mechanisms of defense against brood diseases and parasitic mites. Studies have shown that Apis mellifera colonies remove worker brood infested with Varroa jacobsoni mites from the nest (hygienic behavior), and groom the mites off other adult bees, but to a limited extent compared to the original host of V. jacobsoni, A. cerana. Research is reviewed on hygienic and grooming behaviors with respect to their potential as mechanisms of resistance to V. jacobsoni. Studies related to hygienic behavior include the removal of experimentally infested and naturally infested brood, measurements of heritability, the uncapping and recapping of cells containing infested pupae, and the detection of infested brood. Studies on grooming include the process by which a groomer detects and damages a mite found on itself or on another adult bee, how the behavior is quantified, and problems with these methods of quantification. Finally, unresolved questions concerning grooming and the effects of hygienic and non-hygienic behaviors on limiting the population growth of V. jacobsoni are discussed.
Timing of Mating Flights of Neotropical African and European Honey Bee Queens and Drones (Hymenoptera: Apidae) in Eastern Venezuela
(University of Kansas Natural History Museum, 1999) Otis, Gard W.; Taylor, Orley R., Jr.; Spivak, Marla; Winston, Mark L.; Katz, Susan J.; Kukuk, Penelope F.
Mating flight characteristics of reproductives of neotropica lAfrican and European honey bees (Apis mellifera L.) were studied in eastern Venezuela. For queens, mean exit times of all flights and midpoints of mating flights differed significantly between the subspecies, with the midpoints of mating flights by European queens occurring an average of 47 min earlier in the afternoon. Queens of the two subspecies did not differ in other aspects of their orientation and mating flights. Flights of European drones also occurred significantly earlier than those of neotropical African drones. Mating flights of queens and drones of the same subspecies were approximately synchronous. These results are sufficient to explain the weak positive assortative mating that has been reported previously.
Olfactory and behavioral response thresholds to odors of diseased brood differ between hygienic and non-hygienic honey bees (Apis mellifera L.)
(2001) Masterman, R.; Ross, R.; Mesce, K.; Spivak, M.
Through the use of proboscis-extension reflex conditioning, we demonstrate that honey bees ( Apis mellifera L.) bred for hygienic behavior (a behavioral mechanism of disease resistance) are able to discriminate between odors of healthy and diseased brood at a lower stimulus level than bees from a non-hygienic line. Electroantennogram recordings confirmed that hygienic bees exhibit increased olfactory sensitivity to low concentrations of the odor of chalkbrood infected pupae (a fungal disease caused by Ascosphaera apis ). Three-week-old hygienic bees were able to discriminate between the brood odors significantly better than three-week old non-hygienic bees. However, the differential performance in brood odor discrimination was primarily genetically based, not a direct result of age, experience, or the temporary behavioral state of the bee. Lower stimulus thresholds for both the olfactory and behavioral responses of hygienic bees may facilitate their ability to detect, uncap and remove diseased brood rapidly from the nest. In contrast, non-hygienic bees, possessing higher response thresholds, may not be able to detect diseased brood as easily. Our results provide an example of how physiological and behavioral differences between the hygienic and non-hygienic honey bee lines, operating at the level of the individual, could produce colony-specific behavioral phenotypes.
Resistance to American foulbrood disease by honey bee colonies Apis mellifera bred for hygienic behavior
(2001) Spivak, Marla; Reuter, Gary S.
Honey bee colonies, selected for hygienic behavior on the basis of a freeze-killed brood assay, demonstrated resistance to American foulbrood disease. Over two summers in 1998 and 1999, 18 hygienic and 18 non-hygienic colonies containing instrumentally inseminated queens were challenged with comb sections containing spores of the bacterium Paenibacillus larvae subsp. larvae that causes the disease. The strain of bacterium was demonstrated to be resistant to oxytetracycline antibiotic. Seven (39%) hygienic colonies developed clinical symptoms of the disease but five of these recovered (had no visible symptoms) leaving two colonies (11%) with clinical symptoms. In contrast, 100% of the non-hygienic colonies that were challenged developed clinical symptoms, and only one recovered. All non-hygienic colonies had symptoms of naturally occurring chalkbrood disease (Ascosphaera apis) throughout both summers. In contrast 33% of the hygienic colonies developed clinical symptoms of chalkbrood after they were challenged with American foulbrood, but all recovered. The diseased non-hygienic colonies produced significantly less honey than the hygienic colonies.
Seven suggestive quantitative trait loci influence hygienic behavior of honey bees
(Springer, 2002) Lapidge, Keryn L.; Oldroyd, Benjamin P.; Spivak, Marla
In 1964, Walter Rothenbuhler proposed a twogene model to explain phenotypic variance in the remarkable behavior in which honey bee workers remove dead brood from their colonies. Rothenbuhler’s model proposed that one locus controls the uncapping of brood cells containing dead pupae, while a second controls the removal of the cell contents. We show here, through molecular techniques and quantitative trait loci (QTL) linkage mapping, that the genetic basis of hygienic behavior is more complex, and that many genes are likely to contribute to the behavior. In our cross, we detected seven suggestive QTLs associated with hygienic behavior. Each detected QTL controlled only 9–15% of the observed phenotypic variance in the character.
A Sustainable Approach to Controlling Honey Bee Diseases and Varroa Mites
(USDA, 2005) Spivak, Marla; Reuter, Gary
A multifactorial study of the resistance of honeybees Apis mellifera to the mite Varroa destructor over one year in Mexico
(2005) Mondragon, Luis; Spivak, Marla; Vandame, Remy
A one year study was conducted to evaluate the population growth of three kinds of honey bee colonies and Varroa destructor mites in Mexico, and to estimate the relative contributions of three resistance mechanisms of the bees: hygienic behavior, grooming behavior, and reproductive ability of the parasite. Very significant changes over the year were observed in the number of mated female offspring produced per mother mite (Wr), mite fertility and mutilation of V. destructor. These changes were correlated to the total number of mites per colony. A factorial analysis showed that two mechanisms explained the variation in the amount of mites per colony: Wr (r2 = 0.73) and proportion of mutilated mites (r2 = 0.51). A multi-factorial model including these two mechanisms was significant (r2 = 0.97). The mite fecundity and the hygienic behavior could not explain the population changes of the mite, and the different kinds of bees showed no differences in the expression of the resistance mechanisms.
The relationship between hygienic behavior and suppression of mite reproduction as honey bee (Apis mellifera) mechanisms of resistance to Varroa destructor
(2006) Ibrahim, Abdullah; Spivak, Marla
We compared the mechanisms of resistance to Varroa destructor displayed by bees bred for Suppression of Mite Reproduction (SMR) and hygienic behavior (HYG). Mites from SMR and HYG source colonies were introduced into recently sealed SMR and HYG worker brood, and the infested pupae were placed either into recipient colonies or into an incubator. SMR colonies removed significantly more miteinfested pupae than the HYG colonies. The reproductive success (fertility and number of viable female offspring) of mites from both sources on pupae not removed by bees was significantly lower in SMR colonies. Within the incubator, the reproductive success of mites was also lower on SMR worker pupae, and lowest when mites from SMR colonies were introduced on SMR brood. Our findings indicated that bees bred for SMR express hygienic behavior; adult bees selectively remove pupae infested with mites. In addition, there is an effect of SMR pupae that reduces mite reproductive success that requires further investigation.
Field trial of honey bee colonies bred for mechanisms of resistance against Varroa destructor
(2007) Ibrahim, Abdullah; Reuter, Gary S.; Spivak, Marla
We compared colonies selectively bred for both hygienic behavior and Suppression of Mite Reproduction (HYG/SMR) with colonies bred solely for hygienic behavior (HYG) and unselected control colonies. Colonies were evaluated for strength, brood viability, removal of freeze-killed brood, honey production, mite loads on adult bees and within worker brood, and mite reproductive success on worker brood for two years in two locations. By autumn in both years, the HYG/SMR colonies had significantly fewer mites on adult bees and in worker brood compared to the control colonies, and the HYG colonies had intermediate mite populations. Contrary to expectation, there were no differences among the lines in mite reproductive success. Further studies are required to determine if the genes and neural mechanisms that regulate the SMR trait are the same or different from those regulating hygienic behavior.
Hygienic behaviors of honey bees in response to brood experimentally pin-killed or infected with Ascosphaera apis
(2010) Palacio, María Alejandra; Rodriguez, Edgardo; Goncalves, Lionel; Bedascarrasbure, Enrique; Spivak, Marla
Hygienic behavior of honeybees involves inspection, uncapping and removal of diseased and dead brood from the colony. The objective of this work was to study the activities involved in hygienic behavior of individually tagged bees from selected hygienic (H) and non-hygienic (NH) colonies in the presence of chalkbrood infected brood (Ascosphaera apis) or pin-killed brood. No significant difference was detected in the age of bees inspecting, uncapping or removing brood in H and NH colonies; the median age was 15 days for all activities. The percentage of bees that performed these activities was significantly higher in H colonies. In NH colonies the bees that performed this behavior were more persistent but bees in H colonies were more efficient in the removal of the chalkbrood mummies. H colonies began uncapping more rapidly in response to the stimulus of dead brood independent of the method used to kill it. H and NH bees took the same amount of time to remove the mummies once they initiated the uncapping process but NH colonies took longer to remove pin-killed brood. These findings confirm previous behavioral studies on the activities of hygienic and non-hygienic bees toward freeze-killed brood, but this is the first time the entire process from inspection to removal was focused on individual cells containing actual diseased brood.
Propolis and bee health: the natural history and significance of resin use by honey bees
(2010) Simone-Finstrom, Michael; Spivak, Marla
Social immunity, which describes how individual behaviors of group members effectively reduce disease and parasite transmission at the colony level, is an emerging field in social insect biology. An understudied, but significant behavioral disease resistance mechanism in honey bees is their collection and use of plant resins. Honey bees harvest resins with antimicrobial properties from various plant species and bring them back to the colony where they are then mixed with varying amounts of wax and utilized as propolis. Propolis is an apicultural term for the resins when used by bees within a hive. While numerous studies have investigated the chemical components of propolis that could be used to treat human diseases, there is a lack of information on the importance of propolis in regards to bee health. This review serves to provide a compilation of recent research concerning the behavior of bees in relation to resins and propolis, focusing more on the bees themselves and the potential evolutionary benefits of resin collection. Future research goals are also established in order to create a new focus within the literature on the natural history of resin use among the social insects and role that propolis plays in disease resistance.