• 1
    Rosenkranz P, Aumeier P and Ziegelmann B, Biology and control of Varroa destructor. J Invertebr Pathol 103:96119 (2010).
  • 2
    Aronstein KA, Saldivar E, Vega R, Westmiller S and Douglas AE, How Varroa parasitism affects the immunological and nutritional status of the honey bee, Apis mellifera. Insects 3(3):601615 (2012).
  • 3
    Alaux C, Dantec C, Parrinello H and Le Conte Y, Nutrigenomics in honey bees: Digital gene expression analysis of pollen's nutritive effects on healthy and Varroa-parasitized bees. BMC Genomics 12:496510 (2011).
  • 4
    van Dooremalen C, Gerritsen L, Cornelissen B, van der Steen JJM, van Langevelde F and Blacquière T, Winter survival of individual honey bees and honey bee colonies depends on level of Varroa destructor infestation. PLoS One 7(4):e36285 (2012).
  • 5
    Cox-Foster DL, Conlan S, Holmes EC, Palacios G, Evans JD, Moran NA, et al., A metagenomic survey of microbes in honey bee colony collapse disorder. Science 318:283287 (2007).
  • 6
    van Engelsdorp D, Evans JD, Saegerman C, Mullin C, Haubruge E, Nguyen BK, et al., Colony collapse disorder: A descriptive study. PLoS One doi:10.1371/ps.0006481 (2009).
  • 7
    Dainat B, Evans JD, Chen YP, Gauthier L and Neumann P, Predictive markers of honey bee colony collapse. PLoS One doi:10.1371/ ps.0032151 (2012).
  • 8
    van Engelsdorp D, Hayes Jr J, Underwood RM and Pettis JS, A survey of honey bee colony losses in the United States, fall 2008 to spring 2009. J Apic Res 49:714 (2010).
  • 9
    Lodesani M, Colombo M and Spreafico M, Ineffectiveness of Apistan® treatment against the mite Varroa jacobsoni Oud in several districts of Lombardy (Italy). Apidologie 26:6772 (1995).
  • 10
    Mullin CA, Frazier M, Frazier JL, Ashcraft S, Simonds R, Van engelsdorp D, et al., High levels of miticides and agrochemicals in North American apiaries: Implications for honey bee health. PLoS One Mar 19;5(3):e9754 (2010).
  • 11
    Burley LM, Fell RD and Saacke RG, Survival of honey bee (Hymenoptera: Apidae) spermatozoa incubated at room temperature from drones exposed to miticides. J Econ Entomol 101:10811087 (2008).
  • 12
    Gregorc A, Evans JD, Scharf M and Ellis JD, Gene expression in honey bee (Apis mellifera) larvae exposed to pesticides and Varroa mites (Varroa destructor). J Insect Physiol 58:10421049 (2012).
  • 13
    Boncristiani H, Underwood R, Schwarz R, Evans JD, Pettis J and van Engelsdorp D, Direct effect of acaricides on pathogen loads and gene expression levels in honey bees, Apis mellifera. J Insect Physiol 58:613620 (2012).
  • 14
    Bogdanov S, Imdorf A and Kilchenmann V, Residues in wax and honey after Apilife var treatment. Apidologie 29:513524 (1998).
  • 15
    Floris I, Satta A, Cabras P, Garau VL and Angioni A, Comparison between two thymol formulations in the control of Varroa destructor: effectiveness, persistence, and residues. J Econ Entomol 97:187191 (2004).
  • 16
    Rennich K, Pettis J, van Engelsdorp D, Bozarth R, Eversole H, Roccasecca K, et al., 2011–2012 National Honey Bee Disease & Pest Survey Report. Available: [March 2012].
  • 17
    Adamczyk S, Laäzaro R, Peärez-Arquillueä C, Conchello P and Herrera A, Evaluation of residues of essential oil components in honey after different anti-Varroa treatments. J Agric Food Chem 53:1008510090 (2005).
  • 18
    Nozal MJ, Bernal JL, Jiménez JJ, Gonzalez MJ and Higues M, Extraction of thymol, eucalyptol, menthol and camphor residues from honey and beeswax. Determination by gas chromatography with flame ionization detection. J Chromatogr A 954:207215 (2002).
  • 19
    Gashout AH and Guzmán-Novoa E, Acute toxicity of essential oils and other natural compounds to the parasitic mite, Varroa destructor, and to larval and adult worker honey bees (Apis mellifera). J Apicult Res 48:263269 (2009).
  • 20
    Mattila HR, Otis GW, Daley J and Schulz T, Trials of apiguard, a thymol-based miticide part 2. Non-target effects on honey bees. Am Bee J 140:6870 (2000).
  • 21
    Imdorf A, Kilchenmann V and Bogdanov S, Toxizität von Thymol, Campher, Menthol und Eucalyptol auf Varroa Jacobsoni Oud. und Apis mellifera L. im Labortest. Apidologie 26:2731 (1995).
  • 22
    Aupinel P, Fortini D, Dufour H, Tasei JN, Michaud B, Odoux JF, et al., Improvement of artificial feeding in a standard in vitro method for rearing Apis mellifera larvae. Bull Insect 58:107111 (2005).
  • 23
    Aupinel P, Fortini D, Michaud B, Medrzycki P, Padovani E, Przygoda D, et al., Honey bee brood ring-test: Method for testing pesticide toxicity on honeybee brood in laboratory conditions. Julius-Kühn-Archiv 423:96102 (2009).
  • 24
    Alix A, Chauzat MP, Duchard S, Lewis G, Maus C, Miles MJ, et al., Hazards of Pesticides to Bees: 10th International Symposium of the ICP-BR Bee Protection Group, ed. by Oomen PA and Thompson HM. Julius-Kühn-Archiv, Romania, pp. 1527 (2009).
  • 25
    COLOSS WG 3 Workshop In vitro Larval Rearing Workshop. Available: [November 2011]
  • 26
    Aupinel P, Fortini D, Michaud B, Marolleau F, Tasei JN and Odoux JF, Toxicity of dimethoate and fenoxycarb to honey bee brood (Apis mellifera), using a new in vitro standardized feeding method. Pest Manag Sci 63:10901094 (2007).
  • 27
    Chen YP, Higgins JA and Feldlaufer MF, Quantitative real-time reverse transcription-PCR analysis of deformed wing virus infection in the honeybee (Apis mellifera). Appl Environ Microbiol 71:436441 (2005).
  • 28
    Yang X and Cox-Foster DL, Impact of an ectoparasite on the immunity and pathology of an invertebrate: Evidence for host immunosuppression and viral amplification. Proc Natl Acad Sci U S A 102:74707475 (2005).
  • 29
    Koywiwattrakul P, Thompson GJ, Sitthipraneed S, Oldroyd BP and Maleszka R, Effects of carbon dioxide narcosis on ovary activation and gene expression in worker honeybees, Apis mellifera. J Insect Sci 18:536 (2005).
  • 30
    Zou Z, Lopez DL, Kanost MR, Evans JD and Jiang H, Comparative analysis of serine protease-related genes in the honey bee genome: possible involvement in embryonic development and innate immunity. Insect Mol Biol 15:603614 (2006).
  • 31
    Yang X and Cox-Foster D, Effects of parasitization by Varroa destructor on survivorship and physiological traits of Apis mellifera in correlation with viral incidence and microbial challenge. Parasitology 134:405412 (2007).
  • 32
    Newton K, Peters R and Raftos D, Phenoloxidase and QX disease resistance in Sydney rock oysters (Saccostrea glomerata). Dev Comp Immuno 28:565569 (2004).
  • 33
    Merkx-Jacques M and Bede JC, Influence of diet on the larval beet armyworm, Spodoptera exigua, glucose oxidase activity. J Insect Sci 48:19 (2005).
  • 34
    Lovallo N and Cox-Foster DL, Alteration in FAD-glucose dehydrogenase activity and hemocyte behavior contribute to initial disruption of Manduca sexta immune response to Cotesia congregata parasitoids. J Insect Physiol 45:10371048 (1999).
  • 35
    Habig WH, Pabst MJ and Jakoby WB, Glutathione S-transferases. The first enzymatic step in mercapturic acid formation. J Biol Chem 249:71307139 (1974).
  • 36
    Hendriksma HP, Härtel S and Dewenter IS, Honey bee risk assessment: new approaches for in vitro larvae rearing and data analyses. Methods Ecol Evol 2:509517 (2011).
  • 37
    Fox J, Cox Proportional-Hazards Regression for Survival Data. Available: [February 2002]
  • 38
    Guidugli KR, Piulachs MD, Bellés X, Lourenço AP and Simões ZL, Vitellogenin expression in queen ovaries and in larvae of both sexes of Apis mellifera. Arch Insect Biochem Physiol 59:211218 (2005).
  • 39
    Imdorf A, Bogdanov S, Kilchenmann V and Berger T, The acaricidal effects of essential oils from thyme, salvia and hyssop plants (from left to right) have been tested against Varroa destructor. ALP Sci 495:318 (2006).
  • 40
    Atkins EL and Kellum D, Comparative morphogenic and toxicity studies on the effect of pesticides on honeybee brood. J Apicult Res 25:242255 (1986).
  • 41
    Hummelbrunner LA and Isman MB, Acute, sublethal, antifeedant, and synergistic effects of monoterpenoid essential oil compounds on the tobacco cutworm, Spodoptera litura (Lep., Noctuidae). J Agric Food Chem 49:715720 (2001).
  • 42
    Waliwitiya R, Isman MB, Vernon RS and Riseman A, Insecticidal activity of selected monoterpenoids and rosemary oil to Agriotes obscurus (Coleoptera: Elateridae). J Econ Entomol 98:15601565 (2005).
  • 43
    Karpouhtsis I, Pardali E, Feggou E, Kokkini S, Scouras ZG and Mavragani Tsipidou P, Insecticidal and genotoxic activities of oregano essential oils. J Agric Food Chem 46:11111115 (1998).
  • 44
    Hardstone MC and Scott JG, Is Apis mellifera more sensitive to insecticides than other insects? Pest Manag Sci 66:11711180 (2010).
  • 45
    Suchail S, Guez D and Belzunces LP, Discrepancy between acute and chronic toxicity induced by imidacloprid and its metabolites in Apis mellifera. Environ Toxicol Chem Nov;20(11):24826 (2001).
  • 46
    Dom N, Penninck M, Knapen D and Blust R, Discrepancies in the acute versus chronic toxicity of compounds with a designated narcotic mechanism. Chemosphere 87:742749 (2012).
  • 47
    Priestley CM, Williamson EM, Wafford KA and Sattelle DB, Thymol, a constituent of thyme essential oil, is a positive allosteric modulator of human GABA(A) receptors and a homo-oligomeric GABA receptor from Drosophila melanogaster. Br J Pharmacol 140:13631372. (2003).
  • 48
    Enan EE, Molecular response of Drosophila melanogaster tyramine receptor cascade to plant essential oils. Insect Biochem Mol Biol 35:309321 (2005).
  • 49
    Parnas M, Peters M, Dadon D, Lev S, Vertkin I, Slutsky I, et al., Carvacrol is a novel inhibitor of Drosophila TRPL and mammalian TRPM7 channels. Cell Calcium 45:300309 (2009).
  • 50
    Brouwers EVM, Ebert R and Beetsma J, Behavioural and physiological aspects of nurse bees in relation to the composition of larval food during caste differentiation in the honeybee. J Apicult Res 26:1123 (1987).
  • 51
    Asencot M and Lensky Y, The effect of soluble sugars in stored royal jelly on the differentiation of female honeybee (Apis mellifera L.) larvae to queens. Insect Biochem 18:127133 (1988).
  • 52
    Smodis Skerl MI, Kmecl V and Gregorc A, Exposure to pesticides at sublethal level and their distribution within a honey bee (Apis mellifera) colony. Bull Environ Contam Toxicol 85:125128 (2010).
  • 53
    Rortais A, Arnold G, Halm MP and Touffet-Briens F, Modes of honeybees exposure to systemic insecticides: Estimated amounts of contaminated pollen and nectar consumed by different categories of bees. Apidologie 36:7183 (2005).