References 1. Roeder T (1999) Octopamine in invertebrates. Prog Neurobiol 59: 533–561. 2. Verlinden H, Vleugels R, Marchal E, Badisco L, Pfluger HJ, et al. (2010) The role of octopamine in locusts and other arthropods. J Insect Physiol 56: 854–867. 3. Candy DJ (1978) The regulation of locust flight muscle metabolism by octopamine and other compounds. Insect Biochem 8: 177–181. 4. Mentel T, Duch C, Stypa H, Wegener G, Muller U, et al. (2003) Central modulatory neurons control fuel selection in flight muscle of migratory locust. J Neurosci 23: 1109–1113. 5. Davenport AP, Evans PD (1984) Stress-induced changes in the octopamine levels of insect hemolymph. Insect Biochem 14: 135–143. 6. Stern M (1999) Octopamine in the locust brain: cellular distribution and functional significance in an arousal mechanism. Microsc Res Tech 45: 135–141. 7. Wicher D (2007) Metabolic regulation and behavior: how hunger produces arousal - an insect study. Endocr Metab Immune Disord Drug Targets 7: 304–310. 8. Erber J, Kloppenburg P (1995) The modulatory effects of serotonin and octopamine in the visual system of the honey bee (Apis mellifera L.) I. Behavioral analysis of the motion-sensitive antennal reflex. J Comp Physiol A 176: 111–118. 9. Matheson T (1997) Octopamine modulates the responses and presynaptic inhibition of proprioceptive sensory neurones in the locust Schistocerca gregaria. J Exp Biol 200: 1317–1325. 10. Scheiner R, Plu¨ ckhahn S, O¨ney B, Blenau W, Erber J (2002) Behavioural pharmacology of octopamine, tyramine and dopamine in honey bees. Behav Brain Res 136: 545–553. 11. Widmer A, Hoger U, Meisner S, French AS, Torkkeli PH (2005) Spider peripheral mechanosensory neurons are directly innervated and modulated by octopaminergic efferents. J Neurosci 25: 1588–1598. 12. Zhukovskaya MI, Kapitsky SV (2006) Activity modulation in cockroach sensillum: the role of octopamine. J Insect Physiol 52: 76–86. 13. Dierick HA (2008) Fly fighting: octopamine modulates aggression. Curr Biol 18: R161–163. 14. Hoyer SC, Eckart A, Herrel A, Zars T, Fischer SA, et al. (2008) Octopamine in male aggression of Drosophila. Curr Biol 18: 159–167. 15. Brembs B, Christiansen F, Pfluger HJ, Duch C (2007) Flight initiation and maintenance deficits in flies with genetically altered biogenic amine levels. J Neurosci 27: 11122–11131. 16. Fussnecker BL, Smith BH, Mustard JA (2006) Octopamine and tyramine influence the behavioral profile of locomotor activity in the honey bee (Apis mellifera). J Insect Physiol 52: 1083–1092. 17. Saraswati S, Fox LE, Soll DR, Wu CF (2004) Tyramine and octopamine have opposite effects on the locomotion of Drosophila larvae. J Neurobiol 58: 425–441. 18. Farooqui T, Robinson K, Vaessin H, Smith BH (2003) Modulation of early olfactory processing by an octopaminergic reinforcement pathway in the honeybee. J Neurosci 23: 5370–5380. 19. Hammer M, Menzel R (1998) Multiple sites of associative odor learning as revealed by local brain microinjections of octopamine in honeybees. Learn Mem 5: 146–156. 20. Schwaerzel M, Monastirioti M, Scholz H, Friggi-Grelin F, Birman S, et al. (2003) Dopamine and octopamine differentiate between aversive and appetitive olfactory memories in Drosophila. J Neurosci 23: 10495–10502. 21. Unoki S, Matsumoto Y, Mizunami M (2005) Participation of octopaminergic reward system and dopaminergic punishment system in insect olfactory learning revealed by pharmacological study. Eur J Neurosci 22: 1409–1416. 22. Robinson GE, Heuser LM, Le Conte Y, Lenquette F, Hollingworth RM (1999) Neurochemicals aid bee nestmate recognition. Nature 399: 534–535. 23. Spivak M, Masterman R, Ross R, Mesce KA (2003) Hygienic behavior in the honey bee (Apis mellifera L.) and the modulatory role of octopamine. J Neurobiol 55: 341–354. 24. Schulz DJ, Barron AB, Robinson GE (2002) A role for octopamine in honey bee division of labor. Brain Behav Evol 60: 350–359. 25. Schulz DJ, Robinson GE (1999) Biogenic amines and division of labor in honey bee colonies: behaviorally related changes in the antennal lobes and age-related changes in the mushroom bodies. J Comp Physiol A 184: 481–488. 26. Busch S, Tanimoto H (2010) Cellular configuration of single octopamine neurons in Drosophila. J Comp Neurol 518: 2355–2364. 27. Braunig P, Burrows M, Morris OT (2004) Properties of descending dorsal unpaired median (DUM) neurons of the locust suboesophageal ganglion. Acta Biol Hung 55: 13–19. 28. Dacks AM, Christensen TA, Agricola HJ, Wollweber L, Hildebrand JG (2005) Octopamine-immunoreactive neurons in the brain and subesophageal ganglion of the hawkmoth Manduca sexta. J Comp Neurol 488: 255–268. 29. Konings PN, Vullings HG, Geffard M, Buijs RM, Diederen JH, et al. (1988) Immunocytochemical demonstration of octopamine-immunoreactive cells in the nervous system of Locusta migratoria and Schistocerca gregaria. Cell Tissue Res 251: 371–379. 30. Kreissl S, Eichmuller S, Bicker G, Rapus J, Eckert M (1994) Octopamine-like immunoreactivity in the brain and subesophageal ganglion of the honeybee. J Comp Neurol 348: 583–595. 31. Monastirioti M (1999) Biogenic amine systems in the fruit fly Drosophila melanogaster. Microsc Res Tech 45: 106–121. AmOA1 in the Honey Bee Brain 32. Sinakevitch I, Niwa M, Strausfeld NJ (2005) Octopamine-like immunoreactivity in the honey bee and cockroach: comparable organization in the brain and subesophageal ganglion. J Comp Neurol 488: 233–254. 33. Sinakevitch I, Strausfeld NJ (2006) Comparison of octopamine-like immunore- activity in the brains of the fruit fly and blow fly. J Comp Neurol 494: 460–475. 34. Hammer M (1993) An identified neuron mediates the unconditioned stimulus in associative olfactory learning in honeybees. Nature 366: 59–63. 35. Schro¨ter U, Malun D, Menzel R (2007) Innervation pattern of suboesophageal ventral unpaired median neurones in the honeybee brain. Cell Tissue Res 327: 647–667. 36. Kloppenburg P, Erber J (1995) The Modulatory Effects of serotonin and octopamine in the visual-system of the honey-bee (Apis-Mellifera L) .2. Electrophysiological analysis of motion-sensitive neurons in the lobula. J Comp Physiol A 176: 119–129. 37. Blenau W, Baumann A (2001) Molecular and pharmacological properties of insect biogenic amine receptors: lessons from Drosophila melanogaster and Apis mellifera. Arch Insect Biochem Physiol 48: 13–38. 38. Evans PD, Maqueira B (2005) Insect octopamine receptors: a new classification scheme based on studies of cloned Drosophila G-protein coupled receptors. Invert Neurosci 5: 111–118. 39. Balfanz S, Strunker T, Frings S, Baumann A (2005) A family of octopamine receptors that specifically induce cyclic AMP production or Ca2+ release in Drosophila melanogaster. J Neurochem 93: 440–451. 40. Han KA, Millar NS, Davis RL (1998) A novel octopamine receptor with preferential expression in Drosophila mushroom bodies. J Neurosci 18: 3650–3658. 41. Maqueira B, Chatwin H, Evans PD (2005) Identification and characterization of a novel family of Drosophila beta-adrenergic-like octopamine G-protein coupled receptors. J Neurochem 94: 547–560. 42. Hauser F, Cazzamali G, Williamson M, Blenau W, Grimmelikhuijzen CJ (2006) A review of neurohormone GPCRs present in the fruitfly Drosophila melanogaster and the honey bee Apis mellifera. Prog Neurobiol 80: 1–19. 43. Grohmann L, Blenau W, Erber J, Ebert PR, Strunker T, et al. (2003) Molecular and functional characterization of an octopamine receptor from honeybee (Apis mellifera) brain. J Neurochem 86: 725–735. 44. Lee HG, Seong CS, Kim YC, Davis RL, Han KA (2003) Octopamine receptor OAMB is required for ovulation in Drosophila melanogaster. Dev Biol 264: 179–190. 45. Geffard M, Buijs RM, Seguela P, Pool CW, Le Moal M (1984) First demonstration of highly specific and sensitive antibodies against dopamine. Brain Res 294: 161–165. 46. Sinakevitch I, Geffard M, Pelhate M, Lapied B (1994) Octopamine-like immunoreactivity in the dorsal unpaired median (DUM) neurons innervating the accessory gland of the male cockroach Periplaneta americana. Cell & Tissue Res 276: 15–21. 47. Seguela P, Geffard M, Buijs RM, Le Moal M (1984) Antibodies against gamma- aminobutyric acid: specificity studies and immunocytochemical results. Proc Natl Acad Sci U S A 81: 3888–3892. 48. Sinakevitch I, Douglass JK, Scholtz G, Loesel R, Strausfeld NJ (2003) Conserved and convergent organization in the optic lobes of insects and isopods, with reference to other crustacean taxa. J Comp Neurol 467: 150–172. 49. Sinakevitch IG, Geffard M, Pelhate M, Lapied B (1996) Anatomy and targets of Dorsal Unpaired Median neurones in the Terminal Abdominal Ganglion of the male cockroach Periplaneta americana L. J Comp Neurol 367: 147–163. 50. Brown SM, Strausfeld NJ (2006) Development-dependent and -independent ubiquitin expression in divisions of the cockroach mushroom body. J Comp Neurol 496: 556–571. 51. Sinakevitch I, Farris SM, Strausfeld NJ (2001) Taurine-, aspartate- and glutamate-like immunoreactivity identifies chemically distinct subdivisions of Kenyon cells in the cockroach mushroom body. J Comp Neurol 439: 352–367. 52. Abel R, Rybak J, Menzel R (2001) Structure and response patterns of olfactory interneurons in the honeybee, Apis mellifera. J Comp Neurol 437: 363–383. 53. Flanagan D, Mercer AR (1989) An atlas and 3-D reconstruction of the antennal lobes in the worker honey bee, Apis mellifera (Hymenoptera, Apidae). Int J Insect Morph Embyrol 18: 145–159. 54. Kirschner S, Kleineidam CJ, Zube C, Rybak J, Grunewald B, et al. (2006) Dual olfactory pathway in the honeybee, Apis mellifera. J Comp Neurol 499: 933–952. 55. Arnold G, Masson C, Budharugsa S (1985) Comparative study of the antennal lobes and their afferent pathways in the worker bee and the drone (Apis mellifera). Cell Tissue Res 242: 593–605. 56. Kenyon F (1896) The brain of the bee. A preliminary contribution to the morphology of the nervous system of the Arthropoda. J Comp Neurol 6: 133–210. 57. Farris SM, Abrams AI, Strausfeld NJ (2004) Development and morphology of class II Kenyon cells in the mushroom bodies of the honey bee, Apis mellifera. J Comp Neurol 474: 325–339. 58. Mobbs P (1982) The brain of the honey bee Apis mellifera I. The connections and spatial organization of the mushrrom bodies. Philos Trans R Soc Lond B 298: 309–354. 59. Rybak J, Menzel R (1993) Anatomy of the mushroom bodies in the honey bee brain: the neuronal connections of the alpha-lobe. J Comp Neurol 334: 444–465. PLoS ONE | www.plosone.org 15 January 2011 | Volume 6 | Issue 1 | e14536