Insect Frontiers December 2009 Volume 1 Nomber 2
Transcription
Insect Frontiers December 2009 Volume 1 Nomber 2
December 2009 Volume 1 Nomber 2 Insect Frontiers BPH (brown planthopper)-resistance test of the Bph14transgenic and susceptible wild-type (WT) rice. RI35, resistant parental rice; Kasalath, susceptible WT rice; Ra1–Ra10, Bph14-transgenic T2 lines. (From PNAS 2009 106:2216322168; Abstract No. 9 in this issue. OA article). Insect Frontiers, December 2009 Volume 1 Number 2 Insect Behaviour 1. A polyandrous female moth discriminates against previous mates to gain genetic diversity Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1309-1315 2. Queen replacement in the monogynous ant Aphaenogaster senilis: supernumerary queens as life insurance Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1317-1325 3. Sexual selection for genetic quality: disentangling the roles of male and female behaviour Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1357-1363 4. Policing and dominance behaviour in the parthenogenetic ant Platythyrea punctata Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1427-1431 5. Unexpected absence of behavioural differences between female damselfly colour morphs Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1463-1469 6. Homosexual interactions in bed bugs: alarm pheromones as male recognition signals Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1471-1475 7. Preimaginal and Adult Experience Modulates the Thermal Response Behavior of Ants Current Biology, Volume 19, Issue 22, 1897-1902, 12 November 2009 8. Preferential Ethanol Consumption in Drosophila Models Features of Addiction Current Biology, Volume 19, Issue 24, 2126-2132, 10 December 2009 Insect Biocontrol 9. Identification and characterization of Bph14, a gene conferring resistance to brown planthopper in rice PNAS December 29, 2009 vol. 106 no. 52 22163-22168 Insect Cell Biology 10. A Wolbachia Symbiont in Aedes aegypti Limits Infection with Dengue, Chikungunya, and Plasmodium Cell, Volume 139, Issue 7, 1268-1278, 24 December 2009 11. Increased Cell Bond Tension Governs Cell Sorting at the Drosophila Anteroposterior Compartment Boundary Current Biology, Volume 19, Issue 22, 1950-1955, 29 October 2009 Insect Cellular Metabolisom 12. Schlank, a member of the ceramide synthase family controls growth and body fat in Drosophila The EMBO Journal (2009) 28, 3706 - 3716 Insect Chemoecology 13. Molecular Characterization and Expression Pattern of Two General Odorant Binding Proteins from the Diamondback Moth, Plutella xylostella Journal of Chemical Ecology Volume 35, Number 10 / October, 2009 1188-1196 14. Whiteflies interfere with indirect plant defense against spider mites in Lima bean PNAS December 15, 2009 vol. 106 no. 50 21202-21207 Insect Clock 15. The CRYPTOCHROME Photoreceptor Gates PDF Neuropeptide Signaling to Set Circadian Network Hierarchy in Drosophila Current Biology, Volume 19, Issue 23, 2050-2055, 12 November 2009 Insect Development 16. Function of Drosophila mob2 in photoreceptor morphogenesis Cell and Tissue Research Volume 338, Number 3 / December, 2009 377-389 17. Visualization of Individual Scr mRNAs during Drosophila Embryogenesis Yields Evidence for Transcriptional Bursting 1 Insect Frontiers, December 2009 Volume 1 Number 2 Current Biology, Volume 19, Issue 23, 2037-2042, 19 November 2009 18. Hedgehog, but not Odd skipped, induces segmental grooves in the Drosophila epidermis Development 136, 3875-3880 December 1, 2009. 19. Sanpodo: a context-dependent activator and inhibitor of Notch signaling during asymmetric divisions Development 136, 4089-4098 December 15, 2009. 20. The cadherin Fat2 is required for planar cell polarity in the Drosophila ovary Development 136, 4123-4132 December 15, 2009. 21. Effete-mediated degradation of Cyclin A is essential for the maintenance of germline stem cells in Drosophila Development 136, 4133-4142 December 15, 2009. 22. Drosophila laminins act as key regulators of basement membrane assembly and morphogenesis Development 136, 4165-4176 December 15, 2009. 23. Control in time and space: Tramtrack69 cooperates with Notch and Ecdysone to repress ectopic fate and shape changes during Drosophila egg chamber maturation Development 136, 4187-4197 December 15, 2009. 24. Vasa promotes Drosophila germline stem cell differentiation by activating mei-P26 translation by directly interacting with a (U)-rich motif in its 3′ UTR Genes & Dev. 2009. 23: 2742-2752 Insect Ecology 25. Phase-dependent outbreak dynamics of geometrid moth linked to host plant phenology Proc. R. Soc. B 7 December 2009 vol. 276 no. 1676 4119-4128 Insect Evolution 26. Repeated Co-options of Exoskeleton Formation during Wing-to-Elytron Evolution in Beetles Current Biology, Volume 19, Issue 24, 2057-2065, 10 December 2009 27. EFFECTS OF PARTHENOGENESIS AND GEOGRAPHIC ISOLATION ON FEMALE SEXUAL TRAITS IN A PARASITOID WASP Evolution Volume 63 Issue 12, Pages 3085 – 3096 Published Online: 30 Jul 2009 28. THE TEMPO AND MODE OF EVOLUTION OF TRANSPOSABLE ELEMENTS AS REVEALED BY MOLECULAR PHYLOGENIES RECONSTRUCTED FROM MOSQUITO GENOMES Evolution Volume 63 Issue 12, Pages 3136 – 3146 Published Online: 28 Jul 2009 29. EVOLUTION OF IDENTITY SIGNALS: FREQUENCY-DEPENDENT BENEFITS OF DISTINCTIVE PHENOTYPES USED FOR INDIVIDUAL RECOGNITION Evolution Volume 63 Issue 12, Pages 3106 – 3113 Published Online: 9 Sep 2009 30. Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants PNAS December 15, 2009 vol. 106 no. 50 21236-21241 31. Ejaculate components delay reproductive senescence while elevating female reproductive rate in an insect PNAS December 22, 2009 vol. 106 no. 51 21743-21747 Insect Genetics 32. Little Effect of the tan Locus on Pigmentation in Female Hybrids between Drosophila santomea and D. melanogaster Cell, Volume 139, Issue 6, 1180-1188, 11 December 2009 33. Evolution of the tan Locus Contributed to Pigment Loss in Drosophila santomea: A Response to Matute et al. Cell, Volume 139, Issue 6, 1189-1196, 11 December 2009 34. X-Linked Variation in Immune Response in Drosophila melanogaster Genetics, Vol. 183, 1477-1491, December 2009 35. Altered Heterochromatin Binding by a Hybrid Sterility Protein in Drosophila Sibling Species Science 11 December 2009 Vol. 326. no. 5959, pp. 1538 – 1541 2 Insect Frontiers, December 2009 Volume 1 Number 2 Insect Hearing 36. The cost of assuming the life history of a host: acoustic startle in the parasitoid fly Ormia ochracea Journal of Experimental Biology 212, 4056-4064 (2009) 37. Moths are not silent, but whisper ultrasonic courtship songs Journal of Experimental Biology 212, 4072-4078 (2009) 38. The next step in cicada audition: measuring pico-mechanics in the cicada's ear Journal of Experimental Biology 212, 4079-4083 (2009) 39. Acoustical mimicry in a predatory social parasite of ants Journal of Experimental Biology 212, 4084-4090 (2009) 40. Evaluation of amplitude in male song: female waxmoths respond to fortissimo notes Journal of Experimental Biology 212, 4091-4100 (2009) Insect Learning 41. Interspecific and intersexual learning rate differences in four butterfly species Journal of Experimental Biology 212, 3810-3816 (2009) 42. Thermal learning in the honeybee, Apis mellifera Journal of Experimental Biology 212, 3928-3934 (2009) 43. Structural and proteomic analyses reveal regional brain differences during honeybee aging Journal of Experimental Biology 212, 4027-4032 (2009) 44. Olfactory conditioning of the sting extension reflex in honeybees: Memory dependence on trial number, interstimulus interval, intertrial interval, and protein synthesis Learn. Mem. 2009. 16: 761-765 45. Social learning about egg-laying substrates in fruitflies Proc. R. Soc. B 22 December 2009 vol. 276 no. 1677 4323-4328 Insect Metamorphosis 46. MicroRNA-dependent metamorphosis in hemimetabolan insects PNAS December 22, 2009 vol. 106 no. 51 21678-21682 47. The Insect Neuropeptide PTTH Activates Receptor Tyrosine Kinase Torso to Initiate Metamorphosis Science 4 December 2009: Vol. 326. no. 5958, pp. 1403 – 1405 Insect Molecular Biology 48. Functional involvement of Tudor and dPRMT5 in the piRNA processing pathway in Drosophila germlines The EMBO Journal (2009) 28, 3820 - 3831 49. Pretaporter, a Drosophila protein serving as a ligand for Draper in the phagocytosis of apoptotic cells The EMBO Journal (2009) 28, 3868 – 3878 50. Spatial distribution of opsin-encoding mRNAs in the tiered larval retinas of the sunburst diving beetle Thermonectus marmoratus (Coleoptera: Dytiscidae) Journal of Experimental Biology 212, 3781-3794 (2009) 51. The Drosophila DHR96 nuclear receptor binds cholesterol and regulates cholesterol homeostasis Genes & Dev. 2009. 23: 2711-2716 52. A novel histone fold domain-containing protein that replaces TAF6 in Drosophila SAGA is required for SAGA-dependent gene expression Genes & Dev. 2009. 23: 2818-2823 Insect Neuroethology 53. Local and global motion preferences in descending neurons of the fly Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1107-1120 3 Insect Frontiers, December 2009 Volume 1 Number 2 54. A descending contralateral directionally selective movement detector in the praying mantis Tenodera aridifolia Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1131-1139 Insect Neuroscience 55. Ems and Nkx6 are central regulators in dorsoventral patterning of the Drosophila brain Development 136, 3937-3947 December 1, 2009. 56. The target of rapamycin complex 2 controls dendritic tiling of Drosophila sensory neurons through the Tricornered kinase signalling pathway The EMBO Journal (2009) 28, 3879 – 3892 57. Imaging neural activity in worms, flies and mice with improved GCaMP calcium indicators Nature Methods 6, 875 - 881 (2009) Published online: 8 November 2009 58. Leucine-rich repeat transmembrane proteins instruct discrete dendrite targeting in an olfactory map Nature Neuroscience 12, 1542 - 1550 (2009) Published online: 15 November 2009 59. Frequency Transitions in Odor-Evoked Neural Oscillations Neuron, Volume 64, Issue 5, 692-706, 10 December 2009 60. Glycogen synthase kinase-3/Shaggy mediates ethanol-induced excitotoxic cell death of Drosophila olfactory neurons PNAS December 8, 2009 vol. 106 no. 49 20924-20929 61. Peripheral modulation of worker bee responses to queen mandibular pheromone PNAS December 8, 2009 vol. 106 no. 49 20930-20935 Insect Nutrition 62. Amino-acid imbalance explains extension of lifespan by dietary restriction in Drosophila Nature 462, 1061-1064 (24 December 2009) Insect Pharmacology 63. Transduction mechanism(s) of Na-saccharin in the blowfly Protophormia terraenovae: evidence for potassium and calcium conductance involvement Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1141-1151 Insect Photoreception 64. Specialized ommatidia of the polarization-sensitive dorsal rim area in the eye of monarch butterflies have non-functional reflecting tapeta Cell and Tissue Research Volume 338, Number 3 / December, 2009 391-400 Insect Physiology 65. Isoform- and cell-specific function of tyrosine decarboxylase in the Drosophila Malpighian tubule Journal of Experimental Biology 212, 3802-3809 (2009) Insect Sex Determination 66. The origin of a selfish B chromosome triggering paternal sex ratio in the parasitoid wasp Trichogramma kaykai Proc. R. Soc. B 7 December 2009 vol. 276 no. 1676 4149-4154 Insect Society 67. Brood pheromone suppresses physiology of extreme longevity in honeybees (Apis mellifera) Journal of Experimental Biology 212, 3795-3801 (2009) 68. Flexible task allocation and the organization of work in ants Proc. R. Soc. B 22 December 2009 vol. 276 no. 1677 4373-4380 69. Polymorphic social organization in an ant Proc. R. Soc. B 22 December 2009 vol. 276 no. 1677 4423-4431 4 Insect Frontiers, December 2009 Volume 1 Number 2 Insect Vision 70. Visual detection of diminutive floral guides in the bumblebee Bombus terrestris and in the honeybee Apis mellifera Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1121-1130 Insect Review 71. Are Bigger Brains Better? Current Biology, Volume 19, Issue 21, R995-R1008, 17 November 2009 72. A Review of Ant Cuticular Hydrocarbons Journal of Chemical Ecology Volume 35, Number 10 / October, 2009 1151-1161 73. Courtship learning in Drosophila melanogaster: Diverse plasticity of a reproductive behavior Learn. Mem. 2009. 16: 743-750 74. Circadian rhythms and the evolution of photoperiodic timing in insects Physiological Entomology Volume 34 Issue 4, Pages 301 – 308 Published Online: 19 Nov 2009 5 Insect Frontiers, December 2009 Volume 1 Number 2 Insect Behaviour 1. A polyandrous female moth discriminates against previous mates to gain genetic diversity Jin Xua and Qiao Wanga, a Institute of Natural Resources, Massey University, New Zealand Polyandry is common in many animal species. The hypotheses proposed to explain the evolution of polyandry generally fall into two categories: to gain material benefits and to obtain genetic benefits. Here we report our findings on a polyandrous moth, Ephestia kuehniella (Lepidoptera: Pyralidae). The present study does not support the hypothesis that females gain material benefits from multiple copulations in terms of sperm replenishment, fecundity and longevity, because females that copulated once and females that copulated more than once with the same or different males had similar fertility, fecundity and longevity. Females used three strategies to gain possible genetic benefits: discriminating against previous mates probably via chemical cues, giving 60% chance of paternity to second males, and adjusting their oviposition patterns depending on whether they encountered new or previous mates after the first copulation. For the third strategy, females that encountered new mates after the first copulation saved eggs until the next day to facilitate fertilization of their eggs with the sperm of their new mates. Our findings do not support the hypothesis that polyandry increases offspring viability, because polyandry did not affect egg hatching success, offspring survival or weight. Ephestia kuehniella showed limited ability to disperse, and females laid all their eggs locally, mostly within 2 days, suggesting that offspring live together and that sib copulations are common in this species. Therefore, polyandry is probably a mechanism for reducing sib competition and inbreeding costs in offspring. Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1309-1315 http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6W9W-4XHT4FV3&_user=10&_coverDate=12%2F31%2F2009&_rdoc=1&_fmt=&_orig=search&_sort=d&_docanchor=&view=c&_acct=C000050221 &_version=1&_urlVersion=0&_userid=10&md5=66cbb05a9c470d5aec2758f4759b03ea 2. Queen replacement in the monogynous ant Aphaenogaster senilis: supernumerary queens as life insurance Blandine Chérona, Claudie Doumsa, b, Pierre Fédéricia and Thibaud Monnina a Laboratoire Écologie & Évolution CNRS UMR 7625, Université Pierre et Marie Curie - Paris 6, France b École Pratique des Hautes Études, Paris, France In social groups, the replacement of the breeder may generate conflicts. In eusocial hymenoptera these can occur between queens, between queens and workers, or between workers. We investigated queen replacement in the Mediterranean ant Aphaenogaster senilis, a monogynous (one queen per colony) species that reproduces exclusively by colony fission. We first investigated the potential for worker nepotism by analysing the sociogenetic structure in 31 colonies using microsatellite markers. No polyandry was detected, which eliminates the possibility of worker nepotism. Second, 30 colonies were experimentally orphaned to analyse the production of gynes (unmated, hopeful queens). Workers reared on average two gynes from the previous queen's diploid brood, with the firstborn gyne emerging on average 17 days before the other, supernumerary, gyne(s). The production of only a few gynes severely limits the potential for gyne selection based on quality criteria. Furthermore, such selection would be biased by the asynchrony of gyne emergence which precludes comparing gynes of the same age and maturity. Behavioural observations showed that gynes interacted aggressively and that the firstborn gyne was always dominant. She usually survived while supernumerary gynes seldom did, even though they did not differ in weight. Workers also attacked gynes, and preferentially targeted supernumerary gynes. We propose that supernumerary gynes are produced as insurance in the eventuality of the death of the firstborn gyne, and that the 6 Insect Frontiers, December 2009 Volume 1 Number 2 delay between the emergence of gynes minimizes the risk that aggressive interactions between gynes will lead to all gynes being killed. Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1317-1325 http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6W9W-4XDFDNJ1&_user=10&_coverDate=12%2F31%2F2009&_rdoc=1&_fmt=&_orig=search&_sort=d&_docanchor=&view=c&_acct=C000050221 &_version=1&_urlVersion=0&_userid=10&md5=c02ec861e4282f17e7c91887fffee6a4 3. Sexual selection for genetic quality: disentangling the roles of male and female behaviour Nina Pekkala aMikael Puurtinena and Janne S. Kotiahoa a Centre of Excellence in Evolutionary Research, Department of Biological and Environmental Science, University of Jyväskylä, Finland According to the good genes model of sexual selection, females choose males of good heritable genetic quality to obtain offspring with high fitness. However, better mating success of high-quality males can also be brought about by direct interference competition between males, or simply through elevated activity of high-quality males. We examined the roles of different processes leading to sexual selection for genetic quality in Drosophila montana. We manipulated genetic quality of male flies by inducing mutations with ionizing radiation. We then recorded the effects of inherited heterozygous mutations on several aspects of mating behaviour of males and females in two experiments. We found that mutations reduced the probability of courtship and extended the latency to courtship of the males, suggesting male activity plays a role in selection for genetic quality. However, the effects of mutations on mating success and mating behaviour of the flies were in general weak. No evidence for female mate choice or interference competition between males acting against heritable mutations was found. Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1357-1363 http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6W9W-4XFXSWD1&_user=10&_coverDate=12%2F31%2F2009&_rdoc=12&_fmt=high&_orig=browse&_srch=docinfo(%23toc%236693%232009%23999219993%231571357%23FLA%23display%23Volume)&_cdi=6693&_sort=d&_docanchor=& view=c&_ct=36&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=7310425580e7abc351dd95f61377b259 4. Policing and dominance behaviour in the parthenogenetic ant Platythyrea punctata Elisabeth Brunner1, a, Katrin Kellner, 1, a, and Jürgen Heinzea a Biologie I, University of Regensburg, Germany In the parthenogenetic ant Platythyrea punctata policing behaviour is not expected on relatedness grounds as workers are normally clonemates and thus equally related to all offspring in the colony. Nevertheless, colonies usually contain only a single reproductive and other workers that begin to lay eggs are attacked by their nestmates (‘policing’). We found that those individuals that most actively engaged in policing later themselves had activated ovaries when the old reproductive was removed from the colonies. This suggests that police workers, through attacking and eliminating others, increase their own chances of becoming reproductive themselves. Because regular parthenogenesis leads to a clonal colony structure, individuals are not expected to invest energy in dominance and policing. On the assumption that physical dominance reflects an individual's reproductive potential, aggression among workers might ensure that the most fecund individual becomes the next reproductive, which would benefit the colony as a whole. Furthermore, aggression among nestmates may be maintained in this species despite predominant clonality, because infrequent sex, recombination or the adoption of alien workers may introduce genetic heterogeneity into the colony. Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1427-1431 http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6W9W-4XFF2RR1&_user=10&_coverDate=12%2F31%2F2009&_rdoc=21&_fmt=high&_orig=browse&_srch=docinfo(%23toc%236693%232009%23999219993%231571357%23FLA%23display%23Volume)&_cdi=6693&_sort=d&_docanchor=& view=c&_ct=36&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=20f7005b4a6f76c8827f10e3a779d240 7 Insect Frontiers, December 2009 Volume 1 Number 2 5. Unexpected absence of behavioural differences between female damselfly colour morphs Arne Iserbyt a, and Hans Van Gossuma a Evolutionary Ecology Group, University of Antwerp, Belgium Males are often selected for higher mating rates than females. As a consequence of this sexual conflict, unreceptive females may suffer fitness costs from excessive male sexual harassment. In a variety of vertebrate and invertebrate species, multiple female morphs coexist in natural populations which have been observed to differ in body colour, in behaviour and also in the amount of male harassment received. However, the degree of harassment on a female morph may depend on the frequency and density of males and female morphs in the population. We quantified harassment rate and subsequent refusal behaviour of males and female morphs of the polymorphic damselfly Nehalennia irene. Unexpectedly and contrary to previous work, female morphs received similar amounts of male harassment and showed mostly the same behaviour. We discuss why differences in morph behaviours may be lacking and how this compares to contemporary explanations for the maintenance and evolution of female-limited polymorphisms. Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1463-1469 http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6W9W-4XJ13T62&_user=10&_coverDate=12%2F31%2F2009&_rdoc=26&_fmt=high&_orig=browse&_srch=docinfo(%23toc%236693%232009%23999219993%231571357%23FLA%23display%23Volume)&_cdi=6693&_sort=d&_docanchor=& view=c&_ct=36&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=e1ab49ca435b43f054f00bc38ec339fc 6. Homosexual interactions in bed bugs: alarm pheromones as male recognition signals Camilla Ryne a, a Department of Ecology, Lund University, Sweden Homosexual mounting is a common behaviour in bed bugs as male sexual interest is directed towards any newly fed individual. The only mode of copulation in the common bed bug, Cimex lectularius, is by traumatic insemination, where the male pierces the female abdomen with his needle-like penis. Homosexual mating would result in abdominal injuries in mounted males, as males lack the female counteradaptive spermalege structure. I here show that bed bug alarm pheromones, previously hypothesized to be a predator chemical defence, can be used by newly fed males to signal their sex and reduce the risk of homosexual mating. Mechanical blocking of the male pheromone glands significantly increased homosexual mounting duration compared to control males, while applying male extracts containing mainly alarm pheromone onto male–female mating pairs completely interrupted or shortened mating duration and reduced sperm transfer. Males confined with other males received piercing scars, demonstrating that homosexual mating occurs. The focal males in the all-male confinement experiment had reduced longevity compared to singly held males, but why this reduction in longevity occurred is not clear. Mounted males thus benefit from being able to discharge alarm pheromones, while mounting males consider the alarm signal a major sex identification cue, suggesting that male bed bugs use alarm pheromone communication to avoid homosexual harassment and mounting. Animal Behaviour Volume 78, Issue 6, December 2009, Pages 1471-1475 http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6W9W-4XHT4FV4&_user=10&_coverDate=12%2F31%2F2009&_rdoc=27&_fmt=high&_orig=browse&_srch=docinfo(%23toc%236693%232009%23999219993%231571357%23FLA%23display%23Volume)&_cdi=6693&_sort=d&_docanchor=& view=c&_ct=36&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=d5c945ad04335678e01374833743116b 7. Preimaginal and Adult Experience Modulates the Thermal Response Behavior of Ants Anja Weidenmüller1,, Christina Mayr1, Christoph Johannes Kleineidam1 and Flavio Roces1 8 1 Insect Frontiers, December 2009 Volume 1 Number 2 Behavioral Physiology and Sociobiology, Biozentrum, Am Hubland, Universität Würzburg, D-97074 Würzburg, Germany Colonies of social insects display an amazing degree of flexibility in dealing with longterm and short-term perturbations in their environment. The key organizational element of insect societies is division of labor. Recent literature suggests that interindividual variability in response thresholds plays an important role in the emergence of division of labor among workers (reviewed in [1,2]). Genetic variation can only partly explain the variability among workers. Here we document the effects of both preimaginal and adult thermal experience on the behavioral differentiation of Camponotus rufipes ant workers. We show that preimaginal temperature (22°C or 32°C during pupal stage) affects temperature-response thresholds and temperature preferences of adult brood-tending workers. We further show that brood-carrying experience gathered as adult during several repeated temperature increases modifies thermal behavior. Experienced workers showed a faster transition from first sensing the temperature stimulus to responding with brood translocation. Developmental plasticity of workers provides a colony with flexibility in dealing with thermal variations and constitutes an important mechanism underlying interindividual variability. Adult thermal experience further fine tunes the behavioral response thresholds and reinforces behavioral differentiation among workers. Current Biology, Volume 19, Issue 22, 1897-1902, 12 November 2009 http://www.cell.com/current-biology/abstract/S0960-9822%2809%2901760-6 8. Preferential Ethanol Consumption in Drosophila Models Features of Addiction Anita V. Devineni1, and Ulrike Heberlein2, 1 Program in Neuroscience, University of California, San Francisco, San Francisco, CA 94143-2822, USA 2 Department of Anatomy, University of California, San Francisco, San Francisco, CA 94143-2822, USA Alcohol addiction is a common affliction with a strong genetic component [1]. Although mammalian studies have provided significant insight into the molecular mechanisms underlying ethanol consumption [2], other organisms such as Drosophila melanogaster are better suited for unbiased, forward genetic approaches to identify novel genes. Behavioral responses to ethanol, such as hyperactivity, sedation, and tolerance, are conserved between flies and mammals [3,4], as are the underlying molecular pathways [5,6,7,8,9]. However, few studies have investigated ethanol self-administration in flies [10]. Here we characterize ethanol consumption and preference in Drosophila. Flies prefer to consume ethanol-containing food over regular food, and this preference increases over time. Flies are attracted to the smell of ethanol, which partially mediates ethanol preference, but are averse to its taste. Preference for consuming ethanol is not entirely explained by attraction to either its sensory or caloric properties. We demonstrate that flies can exhibit features of alcohol addiction. First, flies self-administer ethanol to pharmacologically relevant concentrations. Second, flies will overcome an aversive stimulus in order to consume ethanol. Third, flies rapidly return to high levels of ethanol consumption after a period of imposed abstinence. Thus, ethanol preference in Drosophila provides a new model for studying aspects of addiction. Current Biology, Volume 19, Issue 24, 2126-2132, 10 December 2009 http://www.cell.com/current-biology/abstract/S0960-9822(09)01942-3 Insect Biocontrol 9. Identification and characterization of Bph14, a gene conferring resistance to brown planthopper in rice Bo Dua,1, Weilin Zhanga,1, Bingfang Liua, Jing Hua, Zhe Weia, Zhenying Shia, Ruifeng Hea, Lili Zhua, Rongzhi Chena, Bin Hanb and Guangcun Hea,2 [email protected] a Key Laboratory of Ministry of Education for Plant Development Biology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and 9 b Insect Frontiers, December 2009 Volume 1 Number 2 National Center for Gene Research, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, Shanghai 200233, China Planthoppers are highly destructive pests in crop production worldwide. Brown planthopper (BPH) causes the most serious damage of the rice crop globally among all rice pests. Growing resistant varieties is the most effective and environment-friendly strategy for protecting the crop from BPH. More than 19 BPH-resistance genes have been reported and used to various extents in rice breeding and production. In this study, we cloned Bph14, a gene conferring resistance to BPH at seedling and maturity stages of the rice plant, using a map-base cloning approach. We show that Bph14 encodes a coiled-coil, nucleotide-binding, and leucine-rich repeat (CC-NB-LRR) protein. Sequence comparison indicates that Bph14 carries a unique LRR domain that might function in recognition of the BPH insect invasion and activating the defense response. Bph14 is predominantly expressed in vascular bundles, the site of BPH feeding. Expression of Bph14 activates the salicylic acid signaling pathway and induces callose deposition in phloem cells and trypsin inhibitor production after planthopper infestation, thus reducing the feeding, growth rate, and longevity of the BPH insects. Our work provides insights into the molecular mechanisms of rice defense against insects and facilitates the development of resistant varieties to control this devastating insect. PNAS December 29, 2009 vol. 106 no. 52 22163-22168 http://www.pnas.org/content/106/52/22163.abstract?etoc Top Insect Cell Biology 10.A Wolbachia Symbiont in Aedes aegypti Limits Infection with Dengue, Chikungunya, and Plasmodium Luciano A. Moreira1, 2, Iñaki Iturbe-Ormaetxe1, Jason A. Jeffery3, Guangjin Lu3, Alyssa T. Pyke4, Lauren M. Hedges1, Bruno C. Rocha2, Sonja Hall-Mendelin5, Andrew Day5, Markus Riegler1, 6, Leon E. Hugo3, Karyn N. Johnson1, Brian H. Kay3, Elizabeth A. McGraw1, Andrew F. van den Hurk4, 5, Peter A. Ryan3 and Scott L. O'Neill1, 1 School of Biological Sciences, The University of Queensland, Brisbane QLD 4072, Australia 2 René Rachou Research Institute- FIOCRUZ, Belo Horizonte MG, Brazil 3 Queensland Institute of Medical Research, Post Office Royal Brisbane Hospital, Brisbane QLD 4029, Australia 4 Virology, Queensland Health Forensic and Scientific Services, Coopers Plains QLD 4108, Australia 5 School of Chemical and Molecular Biosciences, The University of Queensland, Brisbane QLD 4072, Australia Wolbachia are maternally inherited intracellular bacterial symbionts that are estimated to infect more than 60% of all insect species. While Wolbachia is commonly found in many mosquitoes it is absent from the species that are considered to be of major importance for the transmission of human pathogens. The successful introduction of a life-shortening strain of Wolbachia into the dengue vector Aedes aegypti that halves adult lifespan has recently been reported. Here we show that this same Wolbachia infection also directly inhibits the ability of a range of pathogens to infect this mosquito species. The effect is Wolbachia strain specific and relates to Wolbachia priming of the mosquito innate immune system and potentially competition for limiting cellular resources required for pathogen replication. We suggest that this Wolbachia-mediated pathogen interference may work synergistically with the life-shortening strategy proposed previously to provide a powerful approach for the control of insect transmitted diseases. Cell, Volume 139, Issue 7, 1268-1278, 24 December 2009 http://www.cell.com/abstract/S0092-8674%2809%2901500-1 11.Increased Cell Bond Tension Governs Cell Sorting at the Drosophila Anteroposterior Compartment Boundary 10 Insect Frontiers, December 2009 Volume 1 Number 2 Katharina P. Landsberg1, 3, Reza Farhadifar2, 3, Jonas Ranft2, 3, Daiki Umetsu1, 3, Thomas J. Widmann1, Thomas Bittig2, Amani Said1, Frank Jülicher2, and Christian Dahmann1 1 Max Planck Institute of Molecular Cell Biology and Genetics, Pfotenhauerstrasse 108, 01307 Dresden, Germany 2 Max Planck Institute for the Physics of Complex Systems, Nöthnitzer Strasse 38, 01187 Dresden, Germany Subdividing proliferating tissues into compartments is an evolutionarily conserved strategy of animal development [1,2,3,4,5,6]. Signals across boundaries between compartments can result in local expression of secreted proteins organizing growth and patterning of tissues [1,2,3,4,5,6]. Sharp and straight interfaces between compartments are crucial for stabilizing the position of such organizers and therefore for precise implementation of body plans. Maintaining boundaries in proliferating tissues requires mechanisms to counteract cell rearrangements caused by cell division; however, the nature of such mechanisms remains unclear. Here we quantitatively analyzed cell morphology and the response to the laser ablation of cell bonds in the vicinity of the anteroposterior compartment boundary in developing Drosophila wings. We found that mechanical tension is approximately 2.5-fold increased on cell bonds along this compartment boundary as compared to the remaining tissue. Cell bond tension is decreased in the presence of Y-27632 [7], an inhibitor of Rho-kinase whose main effector is Myosin II [8]. Simulations using a vertex model [9] demonstrate that a 2.5-fold increase in local cell bond tension suffices to guide the rearrangement of cells after cell division to maintain compartment boundaries. Our results provide a physical mechanism in which the local increase in Myosin II-dependent cell bond tension directs cell sorting at compartment boundaries. Current Biology, Volume 19, Issue 22, 1950-1955, 29 October 2009 http://www.cell.com/current-biology/abstract/S0960-9822%2809%2901840-5 Top Insect Cellular Metabolisom 12.Schlank, a member of the ceramide synthase family controls growth and body fat in Drosophila Reinhard Bauer1, André Voelzmann1, Bernadette Breiden2, Ute Schepers2, Hany Farwanah2, Ines Hahn1, Franka Eckardt1, Konrad Sandhoff2 and Michael Hoch1 [email protected]; [email protected] LIMES-Institute, Program Unit Development, Genetics & Molecular Physiology, Laboratory for Molecular Developmental Biology, University of Bonn, Bonn, Germany LIMES-Institute, Program Unit Membrane Biology & Lipid Biochemistry, c/o Kekulé Institute of Organic Chemistry and Biochemistry, University of Bonn, Bonn, Germany Ceramide synthases are highly conserved transmembrane proteins involved in the biosynthesis of sphingolipids, which are essential structural components of eukaryotic membranes and can act as second messengers regulating tissue homeostasis. However, the role of these enzymes in development is poorly understood due to the lack of animal models. We identified schlank as a new Drosophila member of the ceramide synthase family. We demonstrate that schlank is involved in the de novo synthesis of a broad range of ceramides, the key metabolites of sphingolipid biosynthesis. Unexpectedly, schlank mutants also show reduction of storage fat, which is deposited as triacylglyerols in the fat body. We found that schlank can positively regulate fatty acid synthesis by promoting the expression of sterol-responsive element-binding protein (SREBP) and SREBP-target genes. It further prevents lipolysis by downregulating the expression of triacylglycerol lipase. Our results identify schlank as a new regulator of the balance between lipogenesis and lipolysis in Drosophila. Furthermore, our studies of schlank and the mammalian Lass2 family member suggest a novel role for ceramide synthases in regulating body fat metabolism. The EMBO Journal (2009) 28, 3706 - 3716 http://www.nature.com/emboj/journal/v28/n23/abs/emboj2009305a.html 11 Insect Frontiers, December 2009 Volume 1 Number 2 Top Insect Chemoecology 13.Molecular Characterization and Expression Pattern of Two General Odorant Binding Proteins from the Diamondback Moth, Plutella xylostella Zhi-Chun Zhang1, Man-Qun Wang1 , Yao-Bin Lu2 and Guoan Zhang1 [email protected] 1. College of Plant Science and Technology, Huazhong Agricultural University, Wuhan, 430070, China 2. Institute of Plant Protection and Microbiology, Zhejiang Academy of Agricultural Sciences, Hangzhou, 310021, China In the Lepidoptera, odorant signals are thought to be mediated by general odorant binding proteins (GOBPs) in the sensillar lymph surrounding the olfactory receptors. We describe the identification and characterization of two new cDNAs encoding GOBPs from the antennae of the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), a species for which no GOBPs have been identified to date. We focused our investigation on this olfactory protein family by using reverse transcription–polymerase chain reaction strategies. The deduced amino acid sequences of PxylGOBP1 and PxylGOBP2 revealed open reading frames of 168 and 163 amino acids, respectively, with six cysteine residues in conserved positions relative to other known GOBPs. The alignment of the mature PxylGOBPs with other Lepidoptera GOBPs showed high sequence identity (70–80%) with other full-length sequences from GenBank. Sequence identity between PxylGOBP1and PxylGOBP2 was only 50%, suggesting that the two proteins belong to different classes of lepidopteran GOBPs. The expression patterns of the two PxylGOBP genes, with respect to tissue distribution and sex, were further investigated by reverse transcription-polymerase chain reaction (RT-PCR) and real-time PCR. Although the two GOBP genes were expressed only in the antennae of both sexes, reflecting the antennal specificity of GOBPs, the transcription levels of these genes depended on the sex, the age, the mating status, and the genes. Journal of Chemical Ecology Volume 35, Number 10 / October, 2009 1188-1196 http://www.springerlink.com/content/f58804m54772272p/ 14.Whiteflies interfere with indirect plant defense against spider mites in Lima bean Peng-Jun Zhanga, Si-Jun Zhenga, Joop J. A. van Loona, Wilhelm Bolandb, Anja Davidb, Roland Mumma and Marcel Dickea,1 [email protected] a Laboratory of Entomology, Wageningen University, P.O. Box 8031, 6700 EH, Wageningen, The Netherlands; and b Max-Planck-Institute for Chemical Ecology, Hans Knoll Strasse 8, D-07745 Jena, Germany Plants under herbivore attack are able to initiate indirect defense by synthesizing and releasing complex blends of volatiles that attract natural enemies of the herbivore. However, little is known about how plants respond to infestation by multiple herbivores, particularly if these belong to different feeding guilds. Here, we report the interference by a phloem-feeding insect, the whitefly Bemisia tabaci, with indirect plant defenses induced by spider mites (Tetranychus urticae) in Lima bean (Phaseolus lunatus) plants. Additional whitefly infestation of spider-mite infested plants resulted in a reduced attraction of predatory mites (Phytoseiulus persimilis) compared to attraction to plants infested by spider mites only. This interference is shown to result from the reduction in (E)-βocimene emission from plants infested by both spider mites and whiteflies. When using exogenous salicylic acid (SA) application to mimic B. tabaci infestation, we observed similar results in behavioral and chemical analyses. Phytohormone and gene-expression analyses revealed that B. tabaci infestation, as well as SA application, inhibited spider mite-induced jasmonic acid (JA) production and reduced the expression of two JAregulated genes, one of which encodes for the P. lunatus enzyme β-ocimene synthase that catalyzes the synthesis of (E)-β-ocimene. Remarkably, B. tabaci infestation 12 Insect Frontiers, December 2009 Volume 1 Number 2 concurrently inhibited SA production induced by spider mites. We therefore conclude that in dual-infested Lima bean plants the suppression of the JA signaling pathway by whitefly feeding is not due to enhanced SA levels. PNAS December 15, 2009 vol. 106 no. 50 21202-21207 http://www.pnas.org/content/106/50/21202.abstract?etoc Top Insect Clock 15.The CRYPTOCHROME Photoreceptor Gates PDF Neuropeptide Signaling to Set Circadian Network Hierarchy in Drosophila Luoying Zhang1, Bridget C. Lear1, Adam Seluzicki1 and Ravi Allada1, 1 Department of Neurobiology and Physiology, Northwestern University, Evanston, IL 60208, USA Circadian clocks in the brain are organized as coupled oscillators that integrate seasonal cues such as light and temperature to time daily behaviors. In Drosophila, the PIGMENT DISPERSING FACTOR (PDF) neuropeptide-expressing morning (M) and non-PDF evening (E) cells are coupled cell groups important for morning and evening behavior, respectively. Depending on day length, either M cells (short days) or E cells (long days) dictate both the morning and the evening phase, a phenomenon that we term network hierarchy. To examine the role of PDF in light-dark conditions, we examined flies lacking both the PDF receptor (PDFR) and the circadian photoreceptor CRYPTOCHROME (CRY). We found that subsets of E cells exhibit molecular oscillations antiphase to those of wildtype flies, single cry mutants, or single Pdfr mutants, demonstrating a potent role for PDF in light-mediated entrainment, specifically in the absence of CRY. Moreover, we find that the evening behavioral phase is more strongly reset by PDF(+) M cells in the absence of CRY. On the basis of our findings, we propose that CRY can gate PDF signaling to determine behavioral phase and network hierarchy. Current Biology, Volume 19, Issue 23, 2050-2055, 12 November 2009 http://www.cell.com/current-biology/abstract/S0960-9822(09)01922-8 Top Insect Development 16.Function of Drosophila mob2 in photoreceptor morphogenesis Ling-Yu Liu1, Cheng-Han Lin1 and Seng-Sheen Fan1 [email protected] 1Department of Life Science, Tunghai University, No. 181, Sec. 3, Taichung-Kan Road, Taichung, 407, Taiwan, Republic of China The Drosophila photoreceptor is a highly polarized cell; a mature photoreceptor cell in Drosophila contains a photosensitive structure (the rhabdomere) and a supporting membrane (stalk) at its apical membrane. In a screen to isolate genes involved in determining stalk and rhabdomere formation, this study has identified the Drosophila mob2 (Dmob2) gene. Dmob2 belongs to a Mob1/phocein domain protein family whose functions are involved in polarized cell growth and asymmetric cell fate determination in yeast. To study the role of Dmob2 in photoreceptor development, we have raised an antibody against the Dmob2 protein. An immunocytochemical study has shown that Dmob2 is mainly localized in the apical membrane of photoreceptor cells during early development. As development proceeds, Dmob2 is gradually confined to the rhabdomere base of the photoreceptor cells. RNA interference (RNAi) for knockdown Dmob2 expression during eye development impairs rhabdomere formation. Our study further shows that the subcellular localization of phosphorylated Moesin and Crumbs in the developing photoreceptor cell is disrupted in Dmob2 RNAi flies. This work thus reports a novel function of Dmob2 in photoreceptor cell development. 13 Insect Frontiers, December 2009 Volume 1 Number 2 Cell and Tissue Research Volume 338, Number 3 / December, 2009 377-389 http://www.springerlink.com/content/6271281k45685307/ 17.Visualization of Individual Scr mRNAs during Drosophila Embryogenesis Yields Evidence for Transcriptional Bursting Adam Paré1,, Derek Lemons1, Dave Kosman1, William Beaver2, Yoav Freund2 and William McGinnis1, 1 Section of Cell and Developmental Biology, Division of Biology, University of California, San Diego, La Jolla, CA 92093, USA 2 Department of Computer Science and Engineering, University of California, San Diego, La Jolla, CA 92093, USA The detection and counting of transcripts within single cells via fluorescent in situ hybridization (FISH) [1,2,3,4,5,6] has allowed researchers to ask quantitative questions about gene expression at the level of individual cells. This method is often preferable to quantitative RT-PCR [7,8,9], because it does not necessitate destruction of the cells being probed and maintains spatial information that may be of interest. Until now, studies using FISH at single-molecule resolution have only been rigorously carried out in isolated cells (e.g., yeast cells or mammalian cell culture). Here, we describe the detection and counting of transcripts within single cells of fixed, whole-mount Drosophila embryos via a combination of FISH, immunohistochemistry, and image segmentation. Our method takes advantage of inexpensive, long RNA probes detected with antibodies [10,11], and we present novel evidence to show that we can robustly detect single mRNA molecules. We use this method to characterize transcription at the endogenous locus of the Hox gene Sex combs reduced (Scr), by comparing a stably expressing group of cells to a group that only transiently expresses the gene. Our data provide evidence for transcriptional bursting [2,5,12,13,14,15,16], as well for divergent “accumulation” and “maintenance” phases of gene activity at the Scr locus. Current Biology, Volume 19, Issue 23, 2037-2042, 19 November 2009 http://www.cell.com/current-biology/abstract/S0960-9822%2809%2901848-X 18.Hedgehog, but not Odd skipped, induces segmental grooves in the Drosophila epidermis Shai Mulinari and Udo Häcker*[email protected] Department of Experimental Medical Science and Lund Strategic Research Center for Stem Cell Biology and Cell Therapy, Lund University,BMC B13, 22184 Lund, Sweden The formation of segmental grooves during mid embryogenesis in the Drosophila epidermis depends on the specification of a single row of groove cells posteriorly adjacent to cells that express the Hedgehog signal. However, the mechanism of groove formation and the role of the parasegmental organizer, which consists of adjacent rows of hedgehog- and wingless-expressing cells, are not well understood. We report that although groove cells originate from a population of Odd skipped-expressing cells, this pair-rule transcription factor is not required for their specification. We further find that Hedgehog is sufficient to specify groove fate in cells of different origin as late as stage 10, suggesting that Hedgehog induces groove cell fate rather than maintaining a preestablished state. Wingless activity is continuously required in the posterior part of parasegments to antagonize segmental groove formation. Our data support an instructive role for the Wingless/Hedgehog organizer in cellular patterning. Development 136, 3875-3880 December 1, 2009. http://dev.biologists.org/content/136/23/3875.abstract 19.Sanpodo: a context-dependent activator and inhibitor of Notch signaling during asymmetric divisions 14 Insect Frontiers, December 2009 Volume 1 Number 2 A. Burcu Babaoglan1,2,Kate M. O'Connor-Giles3,Hemlata Mistry4, Adam Schickedanz2, Beth A. Wilson2 and James B. Skeath2,* [email protected] 1 Program in Developmental Biology, Washington University School of Medicine, St Louis, MO 63110, USA 2 Department of Genetics, Washington University School of Medicine, St Louis, MO 63110, USA 3 Laboratories of Genetics and Molecular Biology, University of Wisconsin,1525 Linden Drive, Madison, WI 53706, USA 4 Department of Biology, Widener University, Chester, PA 19013, USA Asymmetric cell divisions generate sibling cells of distinct fates (‘A’, ‘B’) and constitute a fundamental mechanism that creates cell-type diversity in multicellular organisms. Antagonistic interactions between the Notch pathway and the intrinsic cell-fate determinant Numb appear to regulate asymmetric divisions in flies and vertebrates. During these divisions, productive Notch signaling requires sanpodo, which encodes a novel transmembrane protein. Here, we demonstrate that Drosophila sanpodo plays a dual role to regulate Notch signaling during asymmetric divisions — amplifying Notch signaling in the absence of Numb in the ‘A’ daughter cell and inhibiting Notch signaling in the presence of Numb in the ‘B’ daughter cell. In so doing, sanpodo ensures the asymmetry in Notch signaling levels necessary for the acquisition of distinct fates by the two daughter cells. These findings answer long-standing questions about the restricted ability of Numb and Sanpodo to inhibit and to promote, respectively, Notch signaling during asymmetric divisions. Development 136, 4089-4098 December 15, 2009. http://dev.biologists.org/content/136/24/4089.abstract 20.The cadherin Fat2 is required for planar cell polarity in the Drosophila ovary Ivana Viktorinová, Tina König*, Karin Schlichting and Christian Dahmann† [email protected] Max Planck Institute of Molecular Cell Biology and Genetics,Pfotenhauerstrasse 108, 01307 Dresden, Germany Planar cell polarity is an important characteristic of many epithelia. In the Drosophila wing, eye and abdomen, establishment of planar cell polarity requires the core planar cell polarity genes and two cadherins, Fat and Dachsous. Drosophila Fat2 is a cadherin related to Fat; however, its role during planar cell polarity has not been studied. Here, we have generated mutations in fat2 and show that Fat2 is required for the planar polarity of actin filament orientation at the basal side of ovarian follicle cells. Defects in actin filament orientation correlate with a failure of egg chambers to elongate during oogenesis. Using a functional fosmid-based fat2-GFP transgene, we show that the distribution of Fat2 protein in follicle cells is planar polarized and that Fat2 localizes where basal actin filaments terminate. Mosaic analysis demonstrates that Fat2 acts nonautonomously in follicle cells, indicating that Fat2 is required for the transmission of polarity information. Our results suggest a principal role for Fat-like cadherins during the establishment of planar cell polarity. Development 136, 4123-4132 December 15, 2009. http://dev.biologists.org/content/136/24/4123.abstract 21.Effete-mediated degradation of Cyclin A is essential for the maintenance of germline stem cells in Drosophila Dongsheng Chen*, Qi Wang*, Haidong Huang*,Laixin Xia, Xiaoyong Jiang, Lijuan Kan, Qinmiao Sun and Dahua Chen† [email protected] State Key Laboratory of Reproductive Biology and State Key Laboratory of Biomembrane and Membrane Biotechnology, Institute of Zoology, Chinese Academy of Sciences,Datun Road, Chaoyang, Beijing 100101, P.R. China 15 Insect Frontiers, December 2009 Volume 1 Number 2 Increasing evidence supports the idea that the regulation of stem cells requires both extrinsic and intrinsic mechanisms. However, much less is known about how intrinsic signals regulate the fate of stem cells. Studies on germline stem cells (GSCs) in the Drosophila ovary have provided novel insights into the regulatory mechanisms of stem cell maintenance. In this study, we demonstrate that a ubiquitin-dependent pathway mediated by the Drosophila eff gene, which encodes the E2 ubiquitin-conjugating enzyme Effete (Eff), plays an essential role in GSC maintenance. We show that Eff both physically and genetically interacts with dAPC2, a key component of the anaphase-promoting complex (APC), which acts as a multisubunit E3 ligase and plays an essential role in targeting mitotic regulators for degradation during exit from mitosis. This interaction indicates that Eff regulates the APC/C-mediated proteolysis pathway in GSCs. Moreover, we show that expression of a stable form of Cyclin A, but not full-length Cyclin A, results in GSC loss. Finally we show that, in common with APC2, Eff is required for the ubiquitylation of Cyclin A, and overexpression of full-length Cyclin A accelerates the loss of GSCs in the eff mutant background. Collectively, our data support the idea that Effete/APC-mediated degradation of Cyclin A is essential for the maintenance of germline stem cells in Drosophila. Given that the regulation of mitotic Cyclins is evolutionarily conserved between flies and mammals, our study also implies that a similar mechanism may be conserved in mammals. Development 136, 4133-4142 December 15, 2009. http://dev.biologists.org/content/136/24/4133.abstract 22.Drosophila laminins act as key regulators of basement membrane assembly and morphogenesis Jose M. Urbano1,*, Catherine N. Torgler1,4,*, Cristina Molnar2,*, Ulrich Tepass3, Ana López-Varea2, Nicholas H. Brown4, Jose F. de Celis2 and Maria D. Martín-Bermudo1,† ([email protected] 1 Centro Andaluz de Biología de Desarrollo (CABD), Univ. Pablo de Olavide-CSIC, Ctra. Utrera Km. 1, 41013 Sevilla, Spain 2 Centro de Biología Molecular ‘Severo Ochoa’, CSIC and Universidad Autónoma de Madrid, Cantoblanco, Madrid 28049, Spain 3 Department of Cell and Systems Biology, University of Toronto,Toronto, Ontario M5S 3G5, Canada 4 The Gurdon Institute and Department of Physiology, Development & Neuroscience, University of Cambridge,Tennis Court Road, Cambridge CB2 1QN, UK Laminins are heterotrimeric molecules found in all basement membranes. In mammals, they have been involved in diverse developmental processes, from gastrulation to tissue maintenance. The Drosophila genome encodes two laminin α chains, one β and one Γ, which form two distinct laminin trimers. So far, only mutations affecting one or other trimer have been analysed. In order to study embryonic development in the complete absence of laminins, we mutated the gene encoding the sole laminin β chain in Drosophila, LanB1, so that no trimers can be made. We show that LanB1 mutant embryos develop until the end of embryogenesis. Electron microscopy analysis of mutant embryos reveals that the basement membranes are absent and the remaining extracellular material appears disorganised and diffuse. Accordingly, abnormal accumulation of major basement membrane components, such as Collagen IV and Perlecan, is observed in mutant tissues. In addition, we show that elimination of LanB1 prevents the normal morphogenesis of most organs and tissues, including the gut, trachea, muscles and nervous system. In spite of the above structural roles for laminins, our results unravel novel functions in cell adhesion, migration and rearrangement. We propose that while an early function of laminins in gastrulation is not conserved in Drosophila and mammals, their function in basement membrane assembly and organogenesis seems to be maintained throughout evolution. Development 136, 4165-4176 December 15, 2009. http://dev.biologists.org/content/136/24/4165.abstract 16 Insect Frontiers, December 2009 Volume 1 Number 2 23.Control in time and space: Tramtrack69 cooperates with Notch and Ecdysone to repress ectopic fate and shape changes during Drosophila egg chamber maturation Michael J. Boyle1 and Celeste A. Berg1,2,* [email protected] 1 Molecular and Cellular Biology Program, University of Washington, Box 355065, Seattle, WA 981955065, USA 2 Department of Genome Sciences, University of Washington,Box 355065, Seattle, WA 98195-5065, USA Organ morphogenesis requires cooperation between cells, which determine their course of action based upon location within a tissue. Just as important, cells must synchronize their activities, which requires awareness of developmental time. To understand how cells coordinate behaviors in time and space, we analyzed Drosophila egg chamber development. We found that the transcription factor Tramtrack69 (TTK69) controls the fates and shapes of all columnar follicle cells by integrating temporal and spatial information, restricting characteristic changes in morphology and expression that occur at stage 10B to appropriate domains. TTK69 is required again later in oogenesis: it controls the volume of the dorsal-appendage (DA) tubes by promoting apical reexpansion and lateral shortening of DA-forming follicle cells. We show that TTK69 and Notch compete to repress each other's expression and that a local Ecdysone signal is required to shift the balance in favor of TTK69. We hypothesize that TTK69 then cooperates with spatially restricted co-factors to define appropriate responses to a globally available (but as yet unidentified) temporal signal that initiates the S10B transformations. Development 136, 4187-4197 December 15, 2009. http://dev.biologists.org/content/136/24/4187.abstract 24.Vasa promotes Drosophila germline stem cell differentiation by activating mei-P26 translation by directly interacting with a (U)-rich motif in its 3′ UTR Niankun Liu, Hong Han and Paul Lasko Department of Biology, Developmental Biology Research Initiative, and Goodman Cancer Centre, McGill University, Montreal, Quebec H3A 1B1, Canada Vasa (Vas) is a DEAD-box RNA-binding protein required in Drosophila at several steps of oogenesis and for primordial germ cell (PGC) specification. Vas associates with eukaryotic initiation factor 5B (eIF5B), and this interaction has been implicated in translational activation of gurken mRNA in the oocyte. Vas is expressed in all ovarian germline cells, and aspects of the vas-null phenotype suggest a function in regulating the balance between germline stem cells (GSCs) and their fate-restricted descendants. We used a biochemical approach to recover Vas-associated mRNAs and obtained mei-P26, whose product represses microRNA activity and promotes GSC differentiation. We found that vas and mei-P26 mutants interact, and that mei-P26 translation is substantially reduced in vas mutant cells. In vitro, Vas protein bound specifically to a (U)-rich motif in the meiP26 3′ untranslated region (UTR), and Vas-dependent regulation of GFP-mei-P26 transgenes in vivo was dependent on the same (U)-rich 3′ UTR domain. The ability of Vas to activate mei-P26 expression in vivo was abrogated by a mutation that greatly reduces its interaction with eIF5B. Taken together, our data support the conclusion that Vas promotes germ cell differentiation by directly activating mei-P26 translation in earlystage committed cells. Genes & Dev. 2009. 23: 2742-2752 http://genesdev.cshlp.org/content/23/23/2742.abstract 17 Insect Frontiers, December 2009 Volume 1 Number 2 Top Insect Ecology 25.Phase-dependent outbreak dynamics of geometrid moth linked to host plant phenology Jane U. Jepsen1,2,*,Snorre B. Hagen1, Stein-Rune Karlsen3 and Rolf A. [email protected] 1. Department of Biology, University of Tromsø, N-9037 Tromsø, Norway 2. Polar Environmental Centre, Norwegian Institute for Nature Research, N-9296 Tromsø, Norway 3. Norut, Northern Research Institute Tromsø, PO Box 6424, N-9294 Tromsø, Norway Climatically driven Moran effects have often been invoked as the most likely cause of regionally synchronized outbreaks of insect herbivores without identifying the exact mechanism. However, the degree of match between host plant and larval phenology is crucial for the growth and survival of many spring-feeding pest insects, suggesting that a phenological match/mismatch-driven Moran effect may act as a synchronizing agent. We analyse the phase-dependent spatial dynamics of defoliation caused by cyclically outbreaking geometrid moths in northern boreal birch forest in Fennoscandia through the most recent massive outbreak (2000–2008). We use satellite-derived time series of the prevalence of moth defoliation and the onset of the growing season for the entire region to investigate the link between the patterns of defoliation and outbreak spread. In addition, we examine whether a phase-dependent coherence in the pattern of spatial synchrony exists between defoliation and onset of the growing season, in order to evaluate if the degree of matching phenology between the moth and their host plant could be the mechanism behind a Moran effect. The strength of regional spatial synchrony in defoliation and the pattern of defoliation spread were both highly phase-dependent. The incipient phase of the outbreak was characterized by high regional synchrony in defoliation and long spread distances, compared with the epidemic and crash phase. Defoliation spread was best described using a two-scale stratified spread model, suggesting that defoliation spread is governed by two processes operating at different spatial scale. The pattern of phase-dependent spatial synchrony was coherent in both defoliation and onset of the growing season. This suggests that the timing of spring phenology plays a role in the large-scale synchronization of birch forest moth outbreaks. Proc. R. Soc. B 7 December 2009 vol. 276 no. 1676 4119-4128 http://rspb.royalsocietypublishing.org/content/276/1676/4119.abstract Top Insect Evolution 26.Repeated Co-options of Exoskeleton Formation during Wing-to-Elytron Evolution in Beetles Yoshinori Tomoyasu1, 4,, Yasuyuki Arakane2, Karl J. Kramer2, 3 and Robin E. Denell1 1 Division of Biology, Kansas State University, Manhattan, KS 66506, USA 2 Department of Biochemistry, Kansas State University, Manhattan, KS 66506, USA 3 Grain Marketing and Production Research Center, Agricultural Research Service, United States Department of Agriculture, 1515 College Avenue, Manhattan, KS 66502, USA The vast diversity in morphology of insect wings provides an excellent model to study morphological evolution. The best-described wing modification is the specification of halteres in Drosophila by a Hox-dependent mechanism, in which a Hox gene affects the expression of genes important for wing development to modify the resulting structure. We have previously shown that highly modified beetle elytra are Hox-free structures despite their divergent morphology, suggesting another mode of evolutionary modification.To understand how elytra have evolved without Hox input, we have analyzed wing development in a coleopteran, Tribolium castaneum. Based on Drosophila mutant phenotypes, we first hypothesized that changes in the wing gene network might 18 Insect Frontiers, December 2009 Volume 1 Number 2 have contributed to elytral evolution. However, we found that the wing gene network defined in Drosophila is largely conserved in Tribolium and is also used to pattern the elytra. Instead, we found evidence that the exoskeleton formation has been co-opted downstream of the conserved wing gene network multiple times. We also show evidence that one of these co-options happened prior to the others, suggesting that repeated cooptions may have strengthened an advantageous trait. In addition, we found that the Tribolium apterous genes are not only essential for exoskeletalization of the elytra but also are required for the proper identity of the hindwing—an unexpected role that we find to be conserved in Drosophila.Our findings suggest that elytral evolution has been achieved by co-opting a beneficial trait several times while conserving the main framework of wing patterning genes. Current Biology, Volume 19, Issue 24, 2057-2065, 10 December 2009 http://www.cell.com/current-biology/abstract/S0960-9822(09)01989-7 27.EFFECTS OF PARTHENOGENESIS AND GEOGRAPHIC ISOLATION ON FEMALE SEXUAL TRAITS IN A PARASITOID WASP Ken Kraaijeveld 1,2 , Padu Franco 1 , Barbara M. Reumer 1 , and Jacques J. M. van Alphen 1,3 1 Animal Ecology, Institute of Biology, University of Leiden, PO Box 9516, 2300 RA Leiden, The Netherlands 2 E-mail: [email protected] 3 UMR 6553 ECOBIO, Université de Rennes I, Campus de Beaulieu, Avenue du Général Leclerc, 35 042 Rennes cedex, France Population divergence in sexual traits is affected by different selection pressures, depending on the mode of reproduction. In allopatric sexual populations, aspects of sexual behavior may diverge due to sexual selection. In parthenogenetic populations, loss-of-function mutations in genes involved in sexual functionality may be selectively neutral or favored by selection. We assess to what extent these processes have contributed to divergence in female sexual traits in the parasitoid wasp Leptopilina clavipes in which some populations are infected with parthenogenesis-inducing Wolbachia bacteria. We find evidence consistent with both hypotheses. Both arrhenotokous males and males derived from thelytokous strains preferred to court females from their own population. This suggests that these populations had already evolved population-specific mating preferences when the latter became parthenogenetic. Thelytokous females did not store sperm efficiently and fertilized very few of their eggs. The nonfertility of thelytokous females was due to mutations in the wasp genome, which must be an effect of mutation accumulation under thelytoky. Divergence in female sexual traits of these two allopatric populations has thus been molded by different forces: independent male/female coevolution while both populations were still sexual, followed by female-only evolution after one population switched to parthenogenesis. Evolution Volume 63 Issue 12, Pages 3085 – 3096 Published Online: 30 Jul 2009 http://www3.interscience.wiley.com/journal/122526547/abstract 28.THE TEMPO AND MODE OF EVOLUTION OF TRANSPOSABLE ELEMENTS AS REVEALED BY MOLECULAR PHYLOGENIES RECONSTRUCTED FROM MOSQUITO GENOMES Claudio J. Struchiner 1,2 , Eduardo Massad 3,4 , Zhijian Tu 5,6 , and José M. C. Ribeiro 7,8 1 ENSP/FIOCRUZ and IMS/UERJ, Av. Brasil, 4365, Rio de Janeiro, Braxil 21040 360 2 E-mail: [email protected] 3 FM/USP, Av. Dr. Arnaldo, 455, São Paulo, Braxil 01246 903 4 E-mail: [email protected] 5 Department of Biochemistry, Virginia Polytechnic Institute and State University, 309 Engel Hall, Blacksburg, Virginia 24061 6 E-mail: [email protected] 7 Laboratory of Malaria and Vector Research, National Institute of Allergy and Infectious Diseases, 12735 Twinbrook Parkway, Room 2E32D, National Institutes of Health, Rockville, Maryland 20852 8 E-mail: [email protected] Although many mathematical models exist predicting the dynamics of transposable elements (TEs), there is a lack of available empirical data to validate these models and inherent assumptions. Genomes can provide a snapshot of several TE families in a single organism, and these could have their demographics inferred by coalescent analysis, allowing for the testing of theories on TE amplification dynamics. Using the available 19 Insect Frontiers, December 2009 Volume 1 Number 2 genomes of the mosquitoes Aedes aegypti and Anopheles gambiae, we indicate that such an approach is feasible. Our analysis follows four steps: (1) mining the two mosquito genomes currently available in search of TE families; (2) fitting, to selected families found in (1), a phylogeny tree under the general time-reversible (GTR) nucleotide substitution model with an uncorrelated lognormal (UCLN) relaxed clock and a nonparametric demographic model; (3) fitting a nonparametric coalescent model to the tree generated in (2); and (4) fitting parametric models motivated by ecological theories to the curve generated in (3). Evolution Volume 63 Issue 12, Pages 3136 – 3146 Published Online: 28 Jul 2009 http://www3.interscience.wiley.com/journal/122523803/abstract 29.EVOLUTION OF IDENTITY SIGNALS: FREQUENCY-DEPENDENT BENEFITS OF DISTINCTIVE PHENOTYPES USED FOR INDIVIDUAL RECOGNITION Michael J. Sheehan 1,2 and Elizabeth A. Tibbetts 1,3 1 Ecology and Evolutionary Biology, University of Michigan, 830 N. University Avenue, Ann Arbor, Michigan 48109 2 E-mail: [email protected] 3 E-mail: [email protected] Identifying broad-scale evolutionary processes that maintain phenotypic polymorphisms has been a major goal of modern evolutionary biology. There are numerous mechanisms, such as negative frequency-dependent selection, that may maintain polymorphisms, although it is unknown which mechanisms are prominent in nature. Traits used for individual recognition are strikingly variable and have evolved independently in numerous lineages, providing an excellent model to investigate which factors maintain ecologically relevant phenotypic polymorphisms. Theoretical models suggest that individuals may benefit by advertising their identities with distinctive, recognizable phenotypes. Here, we test the benefits of advertising one's identity with a distinctive phenotype. We manipulated the appearance of Polistes fuscatus paper wasp groups so that three individuals had the same appearance and one individual had a unique, easily recognizable appearance. We found that individuals with distinctive appearances received less aggression than individuals with nondistinctive appearances. Therefore, individuals benefit by advertising their identity with a unique phenotype. Our results provide a potential mechanism through which negative frequency-dependent selection may maintain the polymorphic identity signals in P. fuscatus. Given that recognition is important for many social interactions, selection for distinctive identity signals may be an underappreciated and widespread mechanism underlying the evolution of phenotypic polymorphisms in social taxa. Evolution Volume 63 Issue 12, Pages 3106 – 3113 Published Online: 9 Sep 2009 http://www3.interscience.wiley.com/journal/122594183/abstract 30.Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants Jacob A. Russella,b,1,2, Corrie S. Moreaua,c,1, Benjamin Goldman-Huertasa, Mikiko Fujiwaraa, David J. Lohmana,d and Naomi E. Piercea [email protected] a Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138; b Department of Biology, Drexel University, Philadelphia, PA 19104; c Department of Zoology, Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605; and d Department of Biology, The City College of The City University of New York, Convent Avenue at 138th Street, New York, NY 10031 Ants are a dominant feature of terrestrial ecosystems, yet we know little about the forces that drive their evolution. Recent findings illustrate that their diets range from herbivorous to predaceous, with “herbivores” feeding primarily on exudates from plants and sap-feeding insects. Persistence on these nitrogen-poor food sources raises the 20 Insect Frontiers, December 2009 Volume 1 Number 2 question of how ants obtain sufficient nutrition. To investigate the potential role of symbiotic microbes, we have surveyed 283 species from 18 of the 21 ant subfamilies using molecular techniques. Our findings uncovered a wealth of bacteria from across the ants. Notable among the surveyed hosts were herbivorous “turtle ants” from the related genera Cephalotes and Procryptocerus (tribe Cephalotini). These commonly harbored bacteria from ant-specific clades within the Burkholderiales, Pseudomonadales, Rhizobiales, Verrucomicrobiales, and Xanthomonadales, and studies of lab-reared Cephalotes varians characterized these microbes as symbiotic residents of ant guts. Although most of these symbionts were confined to turtle ants, bacteria from an antspecific clade of Rhizobiales were more broadly distributed. Statistical analyses revealed a strong relationship between herbivory and the prevalence of Rhizobiales gut symbionts within ant genera. Furthermore, a consideration of the ant phylogeny identified at least five independent origins of symbioses between herbivorous ants and related Rhizobiales. Combined with previous findings and the potential for symbiotic nitrogen fixation, our results strongly support the hypothesis that bacteria have facilitated convergent evolution of herbivory across the ants, further implicating symbiosis as a major force in ant evolution. PNAS December 15, 2009 vol. 106 no. 50 21236-21241 http://www.pnas.org/content/106/50/21236.abstract?etoc 31.Ejaculate components delay reproductive senescence while elevating female reproductive rate in an insect Klaus Reinhardt1, Richard A. Naylor and Michael T. Siva-Jothy [email protected] Department of Animal and Plant Sciences, University of Sheffield, Western Bank, Sheffield S10 2TN, United Kingdom Increased female reproductive rates usually result in accelerated senescence. This correlation provides a link between the evolutionary conflict of the sexes and aging when ejaculate components elevate female reproductive rates at the cost of future reproduction. It is not clear whether this female cost is manifest as shorter lifespan or an earlier onset or a steeper rate of reproductive senescence. It also is unclear whether beneficial ejaculates release females from reproductive trade-offs and, if so, which senescence parameters are affected. We examined these issues in the bedbug, Cimex lectularius, a long-lived insect that shows reduced female lifespan as well as female reproductive senescence at the male-determined mating frequency. We demonstrate experimentally that, independently of the mating frequency, females receiving more ejaculate show increased reproductive rates and enter reproductive senescence later than females receiving less ejaculate. The rate of reproductive senescence did not differ between treatments, and reproductive rates did not predict mortality. The ejaculate effects were consistent in inter- and intra-population crosses, suggesting they have not evolved recently and are not caused by inbreeding. Our results suggest that ejaculate components compensate for the costs of elevated female reproductive rates in bedbugs by delaying the onset of reproductive senescence. Ejaculate components that are beneficial to polyandrous females could have arisen because male traits that protect the ejaculate have positive pleiotropic effects and/or because female counteradaptations to antagonistic male traits exceed the neutralization of those traits. That males influence female reproductive senescence has important consequences for trade-offs between reproduction and longevity and for studies of somatic senescence. PNAS December 22, 2009 vol. 106 no. 51 21743-21747 http://www.pnas.org/content/106/51/21743.abstract?etoc 21 Insect Frontiers, December 2009 Volume 1 Number 2 Top Insect Genetics 32.Little Effect of the tan Locus on Pigmentation in Female Hybrids between Drosophila santomea and D. melanogaster Daniel R. Matute1, Ian A. Butler1 and Jerry A. Coyne1, 1 Department of Ecology and Evolution, The University of Chicago, 1101 E. 57 Street, Chicago, IL 60637, USA Previous work on Drosophila santomea suggested that its absence of abdominal pigmentation, compared to the other darkly pigmented species, is based on mutations in the cis-regulatory region of tan, inactivating the expression of that gene in the abdomen of D. santomea males and females. Our discovery that D. santomea males can produce viable hybrids when mated to D. melanogaster females enables us to use the armamentarium of genetic tools in the latter species to study the genetic basis of this interspecific difference in pigmentation. Hybridization tests using D. melanogaster deficiencies that include tan show no evidence that this locus is involved in the lighter pigmentation of D. santomea females; rather, the pigmentation difference appears to involve at least four other loci in the region. Earlier results implicating tan may have been based on a type of transgenic analysis that can give misleading results about the genes involved in an evolutionary change. Cell, Volume 139, Issue 6, 1180-1188, 11 December 2009 http://www.cell.com/abstract/S0092-8674%2809%2901359-2 33.Evolution of the tan Locus Contributed to Pigment Loss in Drosophila santomea: A Response to Matute et al. Mark Rebeiz1, 5, Margarita Ramos-Womack2, 5, Sangyun Jeong1, 6, Peter Andolfatto3, Thomas Werner1, John True4, David L. Stern2 and Sean B. Carroll1, 1 Howard Hughes Medical Institute and Laboratory of Molecular Biology, University of Wisconsin, 1525 Linden Drive, Madison, WI 53706 USA 2 Howard Hughes Medical Institute and Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544, USA 3 Department of Ecology and Evolutionary Biology and the Lewis-Sigler Institute of Integrative Genomics, Princeton University, Princeton, NJ 08544, USA 4 Department of Ecology and Evolution, Stony Brook University, Stony Brook, NY 11794 USA We have shown previously that the loss of abdominal pigmentation in D. santomea relative to its sister species D. yakuba resulted, in part, from cis-regulatory mutations at the tan locus. Matute et al. claim, based solely upon extrapolation from genetic crosses of D. santomea and D. melanogaster, a much more divergent species, that at least four X chromosome regions but not tan are responsible for pigmentation differences. Here, we provide additional evidence from introgressions of D. yakuba genes into D. santomea that support a causative role for tan in the loss of pigmentation and present analyses that contradict Matute et al.'s claims. We discuss how the choice of parental species and other factors affect the ability to identify loci responsible for species divergence, and we affirm that all of our previously reported results and conclusions stand. Cell, Volume 139, Issue 6, 1189-1196, 11 December 2009 http://www.cell.com/abstract/S0092-8674(09)01377-4 34.X-Linked Variation in Immune Response in Drosophila melanogaster Erin M. Hill-Burns1 and Andrew G. Clark [email protected] Field of Genetics and Development, Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York 14853 22 Insect Frontiers, December 2009 Volume 1 Number 2 This study quantifies the effects of naturally occurring X-linked variation on immune response in Drosophila melanogaster to assess associations between immunity genotypes and innate immune response. We constructed a set of 168 X-chromosomal extraction lines, incorporating X chromosomes from a natural population into co-isogenic autosomal backgrounds, and genotyped the lines at 88 SNPs in 20 X-linked immune genes. We find that genetic variation in many of the genes is associated with immune response phenotypes, including bacterial load and immune gene expression. Many of the associations act in a sex-specific or sexually antagonistic manner, supporting the theory that with the selective pressures facing genes on the X chromosome, sexually antagonistic variation may be more easily maintained. Genetics, Vol. 183, 1477-1491, December 2009 http://www.genetics.org/cgi/content/abstract/183/4/1477?etoc 35.Altered Heterochromatin Binding by a Hybrid Sterility Protein in Drosophila Sibling Species Joshua J. Bayes1,2,* and Harmit S. Malik2,3, [email protected] 1 Molecular and Cellular Biology Program, University of Washington, Seattle, WA 98185, USA. 2 Basic Sciences Division, Fred Hutchinson Cancer Research Center, Seattle, WA 98109, USA. 3 Howard Hughes Medical Institute, Fred Hutchinson Cancer Research Center, Seattle, WA 98109, USA. Hybrid sterility of the heterogametic sex is one of the first postzygotic reproductive barriers to evolve during speciation, yet the molecular basis of hybrid sterility is poorly understood. We show that the hybrid male sterility gene Odysseus-site homeobox (OdsH) encodes a protein that localizes to evolutionarily dynamic loci within heterochromatin and leads to their decondensation. In Drosophila mauritiana x Drosophila simulans male hybrids, OdsH from D. mauritiana (OdsHmau) acts as a sterilizing factor by associating with the heterochromatic Y chromosome of D. simulans, whereas D. simulans OdsH (OdsHsim) does not. Characterization of sterile hybrid testes revealed that OdsH abundance and localization in the premeiotic phases of spermatogenesis differ between species. These results reveal that rapid heterochromatin evolution affects the onset of hybrid sterility. Science 11 December 2009 Vol. 326. no. 5959, pp. 1538 – 1541 http://www.sciencemag.org/cgi/content/abstract/326/5959/1538?sa_campaign=Email/to c/11-December-2009/10.1126/science.1181756 Top Insect Hearing 36. The cost of assuming the life history of a host: acoustic startle in the parasitoid fly Ormia ochracea M. J. Rosen1,*, E. C. Levin2 and R. R. Hoy2 [email protected] 1 Center for Neural Science, New York University, New York, NY 10003, USA 2 Department of Neurobiology and Behavior, Cornell University, Ithaca, NY 14853, USA In the obligatory reproductive dependence of a parasite on its host, the parasite must trade the benefit of ‘outsourcing’ functions like reproduction for the risk of assuming hazards associated with the host. In the present study, we report behavioral adaptations of a parasitic fly, Ormia ochracea, that resemble those of its cricket hosts. Ormia females home in on the male cricket's songs and deposit larvae, which burrow into the cricket, feed and emerge to pupate. Because male crickets call at night, gravid female Ormia in search of hosts are subject to bat predation, in much the same way as female crickets are when responding to male song. We show that Ormia has evolved the same evasive behavior as have crickets: an acoustic startle response to bat-like ultrasound that manifests clearly only during flight. Furthermore, like crickets, Ormia has a sharp 23 Insect Frontiers, December 2009 Volume 1 Number 2 response boundary between the frequencies of song and bat cries, resembling categorical perception first described in the context of human speech. Journal of Experimental Biology 212, 4056-4064 (2009) http://jeb.biologists.org/cgi/content/abstract/212/24/4056?etoc 37. Moths are not silent, but whisper ultrasonic courtship songs R. Nakano1,*, , T. Takanashi2, T. Fujii1, N. Skals3, A. Surlykke4 and Y. Ishikawa1 [email protected] 1 Graduate School of Agricultural and Life Sciences, The University of Tokyo, Bunkyo-ku, Tokyo 1138657, Japan 2 Department of Forest Entomology, Forestry and Forest Products Research Institute, Tsukuba, Ibaraki 305-8687, Japan 3 Vedbendvænget 6, DK-5800 Nyborg, Denmark 4 Center for Sound Communication, Institute of Biology, University of Southern Denmark, DK-5230, Odense M, Denmark Ultrasonic hearing is widespread among moths, but very few moth species have been reported to produce ultrasounds for sexual communication. In those that do, the signals are intense and thus well matched for long distance communication. By contrast, males of the Asian corn borer moth (Crambidae) were recently shown to whisper extremely lowintensity ultrasonic courtship songs close to females. Since low sound levels will prevent eavesdropping by predators, parasites and conspecific rivals, we predicted low intensity ultrasound communication to be widespread among moths. Here we tested 13 species of moths including members of the Noctuidae, Arctiidae, Geometridae and Crambidae. Males of nine species, 70%, produced broadband ultrasound close to females. Peak frequencies ranged from 38 to above 100 kHz. All sounds were of low intensity, 43-76 dB SPL at 1 cm [64±10 dB peSPL (mean ± s.d.), N=9 species]. These quiet and/or hyper-frequency ultrasounds are audible to nearby mates, but inaudible to unintended receivers. Although largely unknown because it is so inconspicuous, acoustic communication using low intensity ultrasound appears to be widespread among hearing moths. Thus, acoustic communication may be the norm rather than the exception. Key words: acoustic communication, courtship behavior, low sound level, moths, ultrasound Journal of Experimental Biology 212, 4072-4078 (2009) http://jeb.biologists.org/cgi/content/abstract/212/24/4072?etoc 38. The next step in cicada audition: measuring pico-mechanics in the cicada's ear J. F. C. Windmill1,*, J. Sueur2 and D. Robert3 [email protected] 1 Centre for Ultrasonic Engineering, Department of Electronic & Electrical Engineering, University of Strathclyde, 204 George Street, Glasgow, G1 1XW, UK 2 Muséum National d'Histoire Naturelle, Département Systématique et Evolution, UMR 7205 CNRS, 45 rue Buffon, F-75231 Paris cedex 05, France 3 School of Biological Sciences, University of Bristol, Woodland Road, Bristol, BS8 1UG, UK Female cicadas use sound when they select a mate from a chorus of singing males. The cicada has a tympanal ear; and the tympanal membrane, and constituent tympanal ridge, act as both acousto-mechanical transducers and frequency filters. The tympanal ridge is physically connected to a large number of mechanoreceptor neurons via a cuticular extension known as the tympanal apodeme. Using microscanning laser Doppler vibrometry, we measured for the first time the in vivo vibrations of the apodeme of female Cicadatra atra in response to the motion of the tympanum driven by sound. These measurements reveal that the nanoscale motion of the tympanal membrane is over a magnitude greater than that of the apodeme. Furthermore, the apodeme acts as an additional mechanical frequency filter, enhancing that of the tympanal ridge, narrowing the frequency band of vibration at the mechanoreceptor neurons to that of the male 24 Insect Frontiers, December 2009 Volume 1 Number 2 calling song. This study enhances our understanding of the mechanical link between the external ear of the cicada and its sensory cells. Journal of Experimental Biology 212, 4079-4083 (2009) http://jeb.biologists.org/cgi/content/abstract/212/24/4079?etoc 39. Acoustical mimicry in a predatory social parasite of ants F. Barbero1,2, S. Bonelli1, J. A. Thomas3, E. Balletto1 and K. Schönrogge2 [email protected] 1 Department of Animal and Human Biology, University of Turin, 10123 Turin, Italy 2 Centre for Ecology and Hydrology, Maclean Building, Wallingford, Oxfordshire, OX10 8BB, UK 3 Department of Zoology, University of Oxford, Tinbergen Building, Oxford, OX1 3PS, UK Rapid, effective communication between colony members is a key attribute that enables ants to live in dominant, fiercely protected societies. Their signals, however, may be mimicked by other insects that coexist as commensals with ants or interact with them as mutualists or social parasites. We consider the role of acoustics in ant communication and its exploitation by social parasites. Social parasitism has been studied mainly in the butterfly genus Maculinea, the final instar larvae of which are host-specific parasites of Myrmica ants, preying either on ant grubs (predatory Maculinea) or being fed by trophallaxis (cuckoo Maculinea). We found similar significant differences between the stridulations of model queen and worker ant castes in both Myrmica sabuleti and Myrmica scabrinodis to that previously reported for Myrmica schencki. However, the sounds made by queens of all three Myrmica species were indistinguishable, and among workers, stridulations did not differ significantly in two of three species-pairs tested. Sounds recorded from the predatory caterpillars and pupae of Maculinea arion had similar or closer patterns to the acoustics of their host Myrmica sabuleti than those previously reported for the cuckoo Maculinea rebeli and its host Myrmica schencki, even though Maculinea rebeli caterpillars live more intimately with their host. We conclude that chemical mimicry enables Maculinea larvae to be accepted as colony members by worker ants, but that caterpillars and pupae of both predatory and cuckoo butterflies employ acoustical mimicry of queen ant calls to elevate their status towards the highest attainable position within their host's social hierarchy. Journal of Experimental Biology 212, 4084-4090 (2009) http://jeb.biologists.org/cgi/content/abstract/212/24/4084?etoc 40. Evaluation of amplitude in male song: female waxmoths respond to fortissimo notes D. Limousin* and M. D. Greenfield [email protected] Institut de recherche sur la biologie de l'insecte (IRBI), CNRS UMR 6035, Université François Rabelais de Tours, Parc de Grandmont, 37200 Tours, France Female evaluation of male signals in the context of sexual selection is often made on the basis of signal energy. Particularly in acoustic species, females may prefer male song that is broadcast at greater amplitude or power. However, song amplitude may be represented by various parameters, and the specific one(s) that are evaluated are not clear. We addressed this problem in an acoustic moth, Achroia grisella (Lepidoptera: Pyralidae), where males attract females with trains of paired ultrasonic pulses. Previous studies showed that females prefer songs that include pulse pairs that have greater mean peak amplitude and that are delivered with greater power (=mean peak amplitudexpulse-pair rate). Here, we report that given male songs of equal acoustic power, females prefer songs in which some pulses attain peak amplitudes that exceed the mean value and that this preference depends largely on the magnitude of amplitude fluctuation. We measured significant variation among males in their degree of amplitude fluctuation, and we note that males that broadcast with lower acoustic power typically show greater relative fluctuations and attain relatively higher amplitude maxima. We discuss the potential role 25 Insect Frontiers, December 2009 Volume 1 Number 2 of multiple integration time constants in female evaluation of mean song amplitude and amplitude maxima. We then consider the possibility that the variation observed in the male population is a response to female choice, but we also indicate that mechanical factors constraining song production may be responsible for such variation. Journal of Experimental Biology 212, 4091-4100 (2009) http://jeb.biologists.org/cgi/content/abstract/212/24/4084?etoc Top Insect Learning 41. Interspecific and intersexual learning rate differences in four butterfly species Ikuo Kandori1,*, Takafumi Yamaki1, Sei-ichi Okuyama2, Noboru Sakamoto2 and Tomoyuki Yokoi3, [email protected] 1 Laboratory of Entomology, Faculty of Agriculture, Kinki University, Japan 2 Itami City Museum of Insects, Japan 3 Laboratory of Insect Ecology, Graduate School of Agriculture, Kyoto University, Japan Learning plays an important role in food acquisition for a wide range of insects and has been demonstrated to be essential during flower foraging in taxa such as bees, parasitoid wasps, butterflies and moths. However, little attention has been focused on differences in floral cue learning abilities among species and sexes. We examined the associative learning of flower colour with nectar in four butterfly species: Idea leuconoe, Argyreus hyperbius, Pieris rapae and Lycaena phlaeas. All butterflies that were trained learned the flower colours associated with food. The flower colour learning rates were significantly higher in I. leuconoe and A. hyperbius than in P. rapae and L. phlaeas. Among the four species examined, the larger and longer-lived species exhibited higher learning rates. Furthermore, female butterflies showed a significantly higher learning rate than males. This study provides the first evidence that learning abilities related to floral cues differ among butterfly species. The adaptive significance of superior learning abilities in the larger and longer-lived butterfly species and in females is discussed. Journal of Experimental Biology 212, 3810-3816 (2009) http://jeb.biologists.org/cgi/content/abstract/212/23/3810?etoc 42. Thermal learning in the honeybee, Apis mellifera Tobin J. Hammer, Curtis Hata and James C. Nieh* [email protected] University of California San Diego, Division of Biological Sciences, Section of Ecology, Behavior, and Evolution, Mail Code 01169500 Gilman Drive, La Jolla, CA 92093-0116, USA Honeybee foragers are exposed to thermal stimuli when collecting food outside and receiving food rewards inside the nest. In both contexts, there is an opportunity for foragers to associate warmth with food rewards. However, honeybee thermal learning is poorly understood. Using an associative learning paradigm (the proboscis extension reflex), we show that honeybees can learn to associate a nectar reward with a heated stimulus applied to the antenna to mimic natural contact with a warm flower or nectaroffering forager. Conditioning with longer inter-trial intervals (ITI) significantly improved learning acquisition. We also trained bees to discriminate between temperatures above (warm) and below (cold) ambient air temperature. Learning acquisition improved by 38% per 10°C increase in absolute stimulus intensity (difference between the rewarded temperature and unrewarded ambient air temperature). However, bees learned positive temperature (warm) significantly better than negative temperature (cold) differences, approximately twice as well for 10°C as compared with a –10°C difference. Thus, thermosensation, a sensory modality that is relatively unexplored in honeybees, could play a role in the acquisition of information from nestmates (social learning) and in 26 Insect Frontiers, December 2009 Volume 1 Number 2 foraging decisions influenced by associations between floral temperature and nectar rewards. Journal of Experimental Biology 212, 3928-3934 (2009) http://jeb.biologists.org/cgi/content/abstract/212/23/3928?etoc 43. Structural and proteomic analyses reveal regional brain differences during honeybee aging F. Wolschin1,2,*, , D. Münch1,* and G. V. Amdam1,2 [email protected] 1 Department of Chemistry, Biotechnology and Food Science, University of Life Sciences, N-1432 Ås, Norway 2 School of Life Sciences, Arizona State University, 85287 Tempe, USA Among insects, learning is particularly well studied in the fruit fly Drosophila melanogaster and the honeybee Apis mellifera. A senescence-dependent decline in classic pavlovian conditioning is demonstrated for both species. To understand how aging affects learning, genetic approaches used with Drosophila can benefit from complementary studies in Apis. Specifically, honeybees have a larger brain size allowing for compartment-specific approaches, and a unique life-history plasticity. They usually perform within-nest tasks early in life (nest bees) and later they collect food (foragers). Senescence of learning performance is a function of the bees' foraging duration but underlying causes are poorly understood. As cognitive aging is commonly associated with structural and biochemical changes in the brain, we hypothesized that brain areas implicated in learning change in synaptic and biochemical composition with increased foraging duration. First, we used synapse-specific immunohistochemistry and proteomics to screen for alterations in the calyx region of the mushroom body, a key structure for memory formation. Using proteomics, we next profiled the central brain, which comprises all higher-order integration centers. We show that, with increased foraging duration, levels of kinases, synaptic- and neuronal growth-related proteins decline in the central brain while the calyx region remains intact both in structure and biochemistry. We suggest that proteome-level changes within major anatomical sites of memory formation other than the calyx region could be central to learning dysfunction. These include large compartments of the central brain, such as the mushroom body's output regions and the antennal lobes. Our data provide novel information toward heterogeneity in the aging insect brain, and demonstrate advantages of the honeybee for invertebrate neurogerontological research. Journal of Experimental Biology 212, 4027-4032 (2009) http://jeb.biologists.org/cgi/content/abstract/212/24/4027 44.Olfactory conditioning of the sting extension reflex in honeybees: Memory dependence on trial number, interstimulus interval, intertrial interval, and protein synthesis Martin Giurfa1,2,4, Eve Fabre1,2,3, Justin Flaven-Pouchon1,2,3, Helga Groll1,2,3, Barbara Oberwallner1,2,3, Vanina Vergoz1,2,3, Edith Roussel1,2 and Jean Christophe Sandoz1,2 [email protected] 1. Université de Toulouse, UPS, Centre de Recherches sur la Cognition Animale, F-31062 Toulouse Cedex 9, France; 2. CNRS, Centre de Recherches sur la Cognition Animale, F-31062 Toulouse Cedex 9, France Harnessed bees learn to associate an odorant with an electric shock so that afterward the odorant alone elicits the sting extension response (SER). We studied the dependency of retention on interstimulus interval (ISI), intertrial interval (ITI), and number of conditioning trials in the framework of olfactory SER conditioning. Forward ISIs (conditioned stimulus [CS] before unconditioned stimulus [US]) supported higher retention than a backward one (US before CS) with an optimum around 3 sec. Spaced trials (ITI 10 min) supported higher retention than massed trials (ITI 1 min) and led to 27 Insect Frontiers, December 2009 Volume 1 Number 2 the formation of a late long-term memory (l-LTM) that depended on protein synthesis. Our results reaffirm olfactory SER conditioning as a reliable tool for the study of learning and memory. Learn. Mem. 2009. 16: 761-765 http://learnmem.cshlp.org/content/16/12/761.abstract 45.Social learning about egg-laying substrates in fruitflies Sachin Sarin and Reuven Dukas* [email protected] Animal Behaviour Group, Department of Psychology, Neuroscience and Behaviour, McMaster University, 1280 Main Street West, Hamilton, ON L8S 4K1, Canada Social learning, defined as learning from other individuals, has had dramatic effects on some species, including humans, in whom it has generated a rich culture. As a first step in examining the evolution of and mechanisms underlying social learning in insects, we tested for social learning in fruitflies (Drosophila melanogaster). Focal females (observers) that experienced novel food together with mated females (models), who had laid eggs on that food, subsequently exhibited a stronger preference for laying eggs on that food over another novel food compared with focal females that experienced the food alone. We observed no social learning, however, when observers experienced food with potentially more ambiguous social information provided by the presence of either virgin models or aggregation pheromone. This first documentation of social learning about egg-laying substrates in fruitflies builds on recent data indicating intricate use of social information by fruitflies and opens up exciting avenues for research on the evolution and neurogenetics of social learning using biology's major model system. Proc. R. Soc. B 22 December 2009 vol. 276 no. 1677 4323-4328 http://rspb.royalsocietypublishing.org/content/276/1677/4323.abstract Top Insect Metamorphosis 46.MicroRNA-dependent metamorphosis in hemimetabolan insects Eva Gomez-Orte and Xavier Belles1 [email protected] Institut de Biologia Evolutiva (Consejo Superior de Investigaciones Científicas and Universitat Pompeu Fabra), Passeig Marítim de la Barceloneta, 37-49, 08003 Barcelona, Spain How does a juvenile insect transform into an adult? This question, which sums up the wonder of insect metamorphosis, has fascinated mankind since ancient times. Modern physiology has established the endocrine basis regulating these transformations, which mainly depend on two hormone types: ecdysteroids, which promote molts, and juvenile hormones, which repress the transformation into the adult stage. The interplay of these two hormones regulates the genes involved in juvenile and adult programs and the shift from one to the other. microRNAs (miRNAs) are small noncoding RNAs, which participate in many biological processes, and we wondered whether they might be also involved in insect metamorphosis. In insects, Dicer-1 ribonuclease transforms miRNA precursors into mature miRNAs. Thus, using systemic RNA interference (RNAi) to silence the expression of Dicer-1 in the hemimetabolan insect Blattella germanica, we depleted miRNA contents in the last instar nymph. This practically inhibited metamorphosis after the next molt, as the resulting specimens showed nymphoid features and were able to molt again. The experiments show that miRNAs play a key role in hemimetabolan metamorphosis, perhaps regulating genes that are juvenile hormone targets. PNAS December 22, 2009 vol. 106 no. 51 21678-21682 http://www.pnas.org/content/106/51/21678.abstract?etoc 28 Insect Frontiers, December 2009 Volume 1 Number 2 47.The Insect Neuropeptide PTTH Activates Receptor Tyrosine Kinase Torso to Initiate Metamorphosis Kim F. Rewitz,1 Naoki Yamanaka,1,2 Lawrence I. Gilbert,3 Michael B. O’Connor1,2,* [email protected] 1 Department of Genetics, Cell Biology, and Development, University of Minnesota, Minneapolis, MN 55455, USA. 2 Howard Hughes Medical Institute, University of Minnesota, Minneapolis, MN 55455, USA. 3 Department of Biology, University of North Carolina, Chapel Hill, NC 27599, USA. Holometabolous insects undergo complete metamorphosis to become sexually mature adults. Metamorphosis is initiated by brain-derived prothoracicotropic hormone (PTTH), which stimulates the production of the molting hormone ecdysone via an incompletely defined signaling pathway. Here we demonstrate that Torso, a receptor tyrosine kinase that regulates embryonic terminal cell fate in Drosophila, is the PTTH receptor. Trunk, the embryonic Torso ligand, is related to PTTH, and ectopic expression of PTTH in the embryo partially rescues trunk mutants. In larvae, torso is expressed specifically in the prothoracic gland (PG), and its loss phenocopies the removal of PTTH. The activation of Torso by PTTH stimulates extracellular signal–regulated kinase (ERK) phosphorylation, and the loss of ERK in the PG phenocopies the loss of PTTH and Torso. We conclude that PTTH initiates metamorphosis by activation of the Torso/ERK pathway. Science 4 December 2009: Vol. 326. no. 5958, pp. 1403 – 1405 http://www.sciencemag.org/cgi/content/abstract/326/5958/1403?sa_campaign=Email/to c/4-December-2009/10.1126/science.1176450 Top Insect Molecular Biology 48.Functional involvement of Tudor and dPRMT5 in the piRNA processing pathway in Drosophila germlines Kazumichi M Nishida1,2, Tomoko N Okada1, Takeshi Kawamura3, Toutai Mituyama4, Yoshinori Kawamura1,2, Sachi Inagaki1, Haidong Huang5, Dahua Chen5, Tatsuhiko Kodama3, Haruhiko Siomi1 and Mikiko C Siomi1,6 [email protected] 1. Department of Molecular Biology, Keio University School of Medicine, Tokyo, Japan 2. Institute of Health Biosciences, University of Tokushima, Tokushima, Japan 3. Research Center for Advanced Science and Technology, University of Tokyo, Tokyo, Japan 4. Computational Biology Research Center (CBRC), National Institute of Advanced Industrial Science and Technology (AIST), Tokyo, Japan 5. Institute of Zoology Chinese Academy of Sciences, Beijing, People's Republic of China 6. JST, CREST, Saitama, Japan In Drosophila, the PIWI proteins, Aubergine (Aub), AGO3, and Piwi are expressed in germlines and function in silencing transposons by associating with PIWI-interacting RNAs (piRNAs). Recent studies show that PIWI proteins contain symmetric dimethylarginines (sDMAs) and that dPRMT5/Capsuleen/DART5 is the modifying enzyme. Here, we show that Tudor (Tud), one of Tud domain-containing proteins, associates with Aub and AGO3, specifically through their sDMA modifications and that these three proteins form heteromeric complexes. piRNA precursor-like molecules are detected in these complexes. The expression levels of Aub and AGO3, along with their degree of sDMA modification, were not changed by tud mutations. However, the population of transposon-derived piRNAs associated with Aub and AGO3 was altered by tud mutations, whereas the total amounts of small RNAs on Aub and AGO3 was increased. Loss of dprmt5 did not change the stability of Aub, but impaired its association with Tud and lowered piRNA association with Aub. Thus, in germline cells, piRNAs are qualitycontrolled by dPRMT5 that modifies PIWI proteins, in tight association with Tud. The EMBO Journal (2009) 28, 3820 - 3831 http://www.nature.com/emboj/journal/v28/n24/abs/emboj2009365a.html 29 Insect Frontiers, December 2009 Volume 1 Number 2 49.Pretaporter, a Drosophila protein serving as a ligand for Draper in the phagocytosis of apoptotic cells Takayuki Kuraishi1,7, Yukiko Nakagawa2, Kaz Nagaosa2, Yumi Hashimoto1, Takashi Ishimoto2, Takeshi Moki2, Yu Fujita1, Hiroshi Nakayama3, Naoshi Dohmae3, Akiko Shiratsuchi1,2,4, Naoko Yamamoto2, Koichi Ueda4, Masamitsu Yamaguchi5, Takeshi Awasaki6 and Yoshinobu Nakanishi1,2,4 [email protected] 1. Graduate School of Medical Science, Kanazawa University, Kanazawa, Ishikawa, Japan 2. Graduate School of Natural Science and Technology, Kanazawa University, Kanazawa, Ishikawa, Japan 3. Biomolecule Characterization Team, RIKEN, Wako, Saitama, Japan 4. Faculty of Pharmaceutical Sciences, Kanazawa University, Kanazawa, Ishikawa, Japan 5. Department of Applied Biology, Kyoto Institute of Technology, Kyoto, Kyoto, Japan 6. Department of Neurobiology, University of Massachusetts Medical School, Worcester, MA, USA Phagocytic removal of cells undergoing apoptosis is necessary for animal development and tissue homeostasis. Draper, a homologue of the Caenorhabditis elegans phagocytosis receptor CED-1, is responsible for the phagocytosis of apoptotic cells in Drosophila, but its ligand presumably present on apoptotic cells remains unknown. An endoplasmic reticulum protein that binds to the extracellular region of Draper was isolated. Loss of this protein, which we name Pretaporter, led to a reduced level of apoptotic cell clearance in embryos, and the overexpression of pretaporter in the mutant flies rescued this defect. Results from genetic analyses suggested that Pretaporter functionally interacts with Draper and the corresponding signal mediators. Pretaporter was exposed at the cell surface after the induction of apoptosis, and cells artificially expressing Pretaporter at their surface became susceptible to Draper-mediated phagocytosis. Finally, the incubation with Pretaporter augmented the tyrosine-phosphorylation of Draper in phagocytic cells. These results collectively suggest that Pretaporter relocates from the endoplasmic reticulum to the cell surface during apoptosis to serve as a ligand for Draper in the phagocytosis of apoptotic cells. The EMBO Journal (2009) 28, 3868 - 3878 http://www.nature.com/emboj/journal/v28/n24/abs/emboj2009343a.html 50. Spatial distribution of opsin-encoding mRNAs in the tiered larval retinas of the sunburst diving beetle Thermonectus marmoratus (Coleoptera: Dytiscidae) Srdjan Maksimovic1, Tiffany A. Cook2 and Elke K. Buschbeck1,* [email protected] 1 Department of Biological Sciences, University of Cincinnati, Cincinnati, Ohio 45221-0006, USA 2 Division of Developmental Biology and Department of Pediatric Ophthalmology, Cincinnati Children's Hospital Medical Center, Cincinnati, Ohio 45229, USA Larvae of the sunburst diving beetle, Thermonectus marmoratus, have a cluster of six stemmata (E1-6) and one eye patch on each side of the head. Each eye has two retinas: a distal retina that is closer to the lens, and a proximal retina that lies directly underneath. The distal retinas of E1 and E2 are made of a dorsal and a ventral stack of at least twelve photoreceptor layers. Could this arrangement be used to compensate for lens chromatic aberration, with shorter wavelengths detected by the distal layers and longer wavelengths by the proximal layers? To answer this question we molecularly identified opsins and their expression patterns in these eyes. We found three opsin-encoding genes. The distal retinas of all six eyes express long-wavelength opsin (TmLW) mRNA, whereas the proximal retinas express ultraviolet opsin (TmUV I) mRNA. In the proximal retinas of E1 and E2, the TmUV I mRNA is expressed only in the dorsal stack. A second ultraviolet opsin mRNA (TmUV II), is expressed in the proximal retinas of E1 and E2 (both stacks). The finding that longer-wavelength opsins are expressed distally to shorter-wavelength opsins makes it unlikely that this retinal arrangement is used to compensate for lens chromatic aberration. In addition, we also described opsin expression patterns in the medial retina of E1 and in the non-tiered retina of the lensless eye patch. To our 30 Insect Frontiers, December 2009 Volume 1 Number 2 knowledge, this is also the first report of multiple UV opsins being expressed in the same stemma. Journal of Experimental Biology 212, 3781-3794 (2009) http://jeb.biologists.org/cgi/content/abstract/212/23/3781?etoc 51.The Drosophila DHR96 nuclear receptor binds cholesterol and regulates cholesterol homeostasis Michael A. Horner1, Keith Pardee2, Suya Liu3, Kirst King-Jones4, Gilles Lajoie3, Aled Edwards2, Henry M. Krause2 and Carl S. Thummel1, 1 Department of Human Genetics, University of Utah School of Medicine, Salt Lake City, Utah 84112, USA; 2 Banting and Best Department of Medical Research, University of Toronto, Toronto Ontario M5G 1L6, Canada; 3 UWO Biological Mass Spectrometry Laboratory, University of Western Ontario, London, Ontario N6G 2V4, Canada; 4 Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, Canada Cholesterol homeostasis is required to maintain normal cellular function and avoid the deleterious effects of hypercholesterolemia. Here we show that the Drosophila DHR96 nuclear receptor binds cholesterol and is required for the coordinate transcriptional response of genes that are regulated by cholesterol and involved in cholesterol uptake, trafficking, and storage. DHR96 mutants die when grown on low levels of cholesterol and accumulate excess cholesterol when maintained on a high-cholesterol diet. The cholesterol accumulation phenotype can be attributed to misregulation of npc1b, an ortholog of the mammalian Niemann-Pick C1-like 1 gene NPC1L1, which is essential for dietary cholesterol uptake. These studies define DHR96 as a central regulator of cholesterol homeostasis. Genes & Dev. 2009. 23: 2711-2716 http://genesdev.cshlp.org/content/23/23/2711.abstract 52.A novel histone fold domain-containing protein that replaces TAF6 in Drosophila SAGA is required for SAGA-dependent gene expression Vikki M. Weake1, Selene K. Swanson1, Arcady Mushegian1,2, Laurence Florens1, Michael P. Washburn1,3, Susan M. Abmayr1,4 and Jerry L. Workman1,[email protected] 1 Stowers Institute for Medical Research, Kansas City, Missouri 64110, USA; 2 Department of Microbiology, Molecular Genetics and Immunology, University of Kansas Medical Center, Kansas City, Kansas 66160, USA; 3 Department of Pathology and Laboratory Medicine, University of Kansas Medical Center, Kansas City, Kansas 66160, USA; 4 Department of Anatomy and Cell Biology, University of Kansas Medical Center, Kansas City, Kansas 66160, USA The histone acetyltransferase complex SAGA is well characterized as a coactivator complex in yeast. In this study of Drosophila SAGA (dSAGA), we describe three novel components that include an ortholog of Spt20, a potential ortholog of Sgf73/ATXN7, and a novel histone fold protein, SAF6 (SAGA factor-like TAF6). SAF6, which binds directly to TAF9, functions analogously in dSAGA to TAF6/TAF6L in the yeast and human SAGA complexes, respectively. Moreover, TAF6 in flies is restricted to TFIID. Mutations in saf6 disrupt SAGA-regulated gene expression without disrupting acetylated or ubiquitinated histone levels. Thus, SAF6 is essential for SAGA coactivator function independent of the enzymatic activities of the complex. Genes & Dev. 2009. 23: 2818-2823 http://genesdev.cshlp.org/content/23/24/2818.abstract?etoc 31 Insect Frontiers, December 2009 Volume 1 Number 2 Top Insect Neuroethology 53.Local and global motion preferences in descending neurons of the fly Adrian Wertz1 , Juergen Haag1 and Alexander Borst1 [email protected] 1 Department of Systems and Computational Neurobiology, Max-Planck-Institute of Neurobiology, Am Klopferspitz 18, 82152 Martinsried, Germany For a moving animal, optic flow is an important source of information about its egomotion. In flies, the processing of optic flow is performed by motion sensitive tangential cells in the lobula plate. Amongst them, cells of the vertical system (VS cells) have receptive fields with similarities to optic flows generated during rotations around different body axes. Their output signals are further processed by pre-motor descending neurons. Here, we investigate the local motion preferences of two descending neurons called descending neurons of the ocellar and vertical system (DNOVS1 and DNOVS2). Using an LED arena subtending 240° × 95° of visual space, we mapped the receptive fields of DNOVS1 and DNOVS2 as well as those of their presynaptic elements, i.e. VS cells 1–10 and V2. The receptive field of DNOVS1 can be predicted in detail from the receptive fields of those VS cells that are most strongly coupled to the cell. The receptive field of DNOVS2 is a combination of V2 and VS cells receptive fields. Predicting the global motion preferences from the receptive field revealed a linear spatial integration in DNOVS1 and a superlinear spatial integration in DNOVS2. In addition, the superlinear integration of V2 output is necessary for DNOVS2 to differentiate between a roll rotation and a lift translation of the fly. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1107-1120 http://www.springerlink.com/content/7p871645856140ul/ 54.A descending contralateral directionally selective movement detector in the praying mantis Tenodera aridifolia Yoshifumi Yamawaki1 and Yoshihiro Toh1 [email protected] 1 Department of Biology, Faculty of Science, Kyushu University, Fukuoka 812-8581, Japan Extracellular recordings were made from a directionally selective neuron in the ventral nerve cord of mantises. The neuron’s preferred direction of motion was forward and upward over the compound eye contralateral to its axon at the cervical connective. The neuron was sensitive to wide-field motion stimuli, resistant to habituation, and showed transient excitation in response to light ON and OFF stimuli. Its responses to drifting gratings depended on the temporal frequency and contrast of the stimulus. These results suggest that the neuron receives input from correlation-type motion detectors. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1131-1139 http://www.springerlink.com/content/t4v72x575n5u7607/ Top Insect Neuroscience 55.Ems and Nkx6 are central regulators in dorsoventral patterning of the Drosophila brain Janina Seibert, Dagmar Volland and Rolf Urbach*[email protected] Institute of Genetics, University of Mainz,D-55099 Mainz, Germany 32 Insect Frontiers, December 2009 Volume 1 Number 2 In central nervous system development, the identity of neural stem cells (neuroblasts) critically depends on the precise spatial patterning of the neuroectoderm in the dorsoventral (DV) axis. Here, we uncover a novel gene regulatory network underlying DV patterning in the Drosophila brain, and show that the cephalic gap gene empty spiracles (ems) and the Nk6 homeobox gene (Nkx6) encode key regulators. The regulatory network implicates novel interactions between these and the evolutionarily conserved homeobox genes ventral nervous system defective (vnd), intermediate neuroblasts defective (ind) and muscle segment homeobox (msh). We show that Msh crossrepressively interacts with Nkx6 to sustain the boundary between dorsal and intermediate neuroectoderm in the tritocerebrum (TC) and deutocerebrum (DC), and that Vnd positively regulates Nkx6 by suppressing Msh. Remarkably, Ems is required to activate Nkx6, ind and msh in the TC and DC, whereas later Nkx6 and Ind act together to repress ems in the intermediate DC. Furthermore, the initially overlapping expression of Ems and Vnd in the ventral/intermediate TC and DC resolves into complementary expression patterns due to cross-repressive interaction. These results indicate that the anteroposterior patterning gene ems controls the expression of DV genes, and vice versa. In addition, in contrast to regulation in the ventral nerve cord, cross-inhibition between homeodomain factors (between Ems and Vnd, and between Nkx6 and Msh) is essential for the establishment and maintenance of discrete DV gene expression domains in the Drosophila brain. This resembles the mutually repressive relationship between pairs of homeodomain proteins that pattern the vertebrate neural tube in the DV axis. Development 136, 3937-3947 December 1, 2009. http://dev.biologists.org/content/136/23/3937.abstract 56.The target of rapamycin complex 2 controls dendritic tiling of Drosophila sensory neurons through the Tricornered kinase signalling pathway Makiko Koike-Kumagai1, Kei-ichiro Yasunaga1, Rei Morikawa1, Takahiro Kanamori1 and Kazuo Emoto1,2,3 [email protected] 1. Neural Morphogenesis Laboratory, National Institute of Genetics, Mishima, Japan 2. Department of Genetics, SOKENDAI, Mishima, Japan 3. PRESTO, Japan Science and Technology Agency, Saitama, Japan To cover the receptive field completely and non-redundantly, neurons of certain functional groups arrange tiling of their dendrites. In Drosophila class IV dendrite arborization (da) neurons, the NDR family kinase Tricornered (Trc) is required for homotypic repulsion of dendrites that facilitates dendritic tiling. We here report that Sin1, Rictor, and target of rapamycin (TOR), components of the TOR complex 2 (TORC2), are required for dendritic tiling of class IV da neurons. Similar to trc mutants, dendrites of sin1 and rictor mutants show inappropriate overlap of the dendritic fields. TORC2 components physically and genetically interact with Trc, consistent with a shared role in regulating dendritic tiling. Moreover, TORC2 is essential for Trc phosphorylation on a residue that is critical for Trc activity in vivo and in vitro. Remarkably, neuronal expression of a dominant active form of Trc rescues the tiling defects in sin1 and rictor mutants. These findings suggest that TORC2 likely acts together with the Trc signalling pathway to regulate the dendritic tiling of class IV da neurons, and thus uncover the first neuronal function of TORC2 in vivo. The EMBO Journal (2009) 28, 3879 - 3892 http://www.nature.com/emboj/journal/v28/n24/abs/emboj2009312a.html 57.Imaging neural activity in worms, flies and mice with improved GCaMP calcium indicators Lin Tian1, S Andrew Hires1, Tianyi Mao1, Daniel Huber1, M Eugenia Chiappe1, Sreekanth H Chalasani2, Leopoldo Petreanu1, Jasper Akerboom1, Sean A McKinney1,4, Eric R Schreiter3, Cornelia I Bargmann2, Vivek Jayaraman1, Karel Svoboda1 & Loren L Looger1 [email protected] 33 Insect Frontiers, December 2009 Volume 1 Number 2 1. 2. 3. 4. Howard Hughes Medical Institute, Janelia Farm Research Campus, Ashburn, Virginia, USA. Howard Hughes Medical Institute, Laboratory of Neural Circuits and Behavior, The Rockefeller University, New York, New York, USA. Department of Chemistry, University of Puerto Rico, Río Piedras, San Juan, Puerto Rico. Present address: The Stowers Institute, Kansas City, Missouri, USA. Genetically encoded calcium indicators (GECIs) can be used to image activity in defined neuronal populations. However, current GECIs produce inferior signals compared to synthetic indicators and recording electrodes, precluding detection of low firing rates. We developed a single-wavelength GCaMP2-based GECI (GCaMP3), with increased baseline fluorescence (3-fold), increased dynamic range (3-fold) and higher affinity for calcium (1.3-fold). We detected GCaMP3 fluorescence changes triggered by single action potentials in pyramidal cell dendrites, with signal-to-noise ratio and photostability substantially better than those of GCaMP2, D3cpVenus and TN-XXL. In Caenorhabditis elegans chemosensory neurons and the Drosophila melanogaster antennal lobe, sensory stimulation–evoked fluorescence responses were significantly enhanced with GCaMP3 (4– 6-fold). In somatosensory and motor cortical neurons in the intact mouse, GCaMP3 detected calcium transients with amplitudes linearly dependent on action potential number. Long-term imaging in the motor cortex of behaving mice revealed large fluorescence changes in imaged neurons over months. Nature Methods 6, 875 - 881 (2009) Published online: 8 November 2009 http://www.nature.com/nmeth/journal/v6/n12/abs/nmeth.1398.html?lang=en 58.Leucine-rich repeat transmembrane proteins instruct discrete dendrite targeting in an olfactory map Weizhe Hong1, Haitao Zhu1,4, Christopher J Potter1, Gabrielle Barsh1, Mitsuhiko Kurusu2,3, Kai Zinn2 & Liqun Luo1 [email protected] 1. 2. 3. 4. Howard Hughes Medical Institute and Department of Biology, Stanford University, Stanford, California, USA. Division of Biology, California Institute of Technology, Pasadena, California, USA. Structural Biology Center, National Institute of Genetics, and Department of Genetics, the Graduate University for Advanced Studies, Mishima, Japan. Present address: Department of Neurodegeneration, Genentech, South San Francisco, California, USA. Olfactory systems utilize discrete neural pathways to process and integrate odorant information. In Drosophila, axons of first-order olfactory receptor neurons (ORNs) and dendrites of second-order projection neurons (PNs) form class-specific synaptic connections at 50 glomeruli. The mechanisms underlying PN dendrite targeting to distinct glomeruli in a three-dimensional discrete neural map are unclear. We found that the leucine-rich repeat (LRR) transmembrane protein Capricious (Caps) was differentially expressed in different classes of PNs. Loss-of-function and gain-of-function studies indicated that Caps instructs the segregation of Caps-positive and Caps-negative PN dendrites to discrete glomerular targets. Moreover, Caps-mediated PN dendrite targeting was independent of presynaptic ORNs and did not involve homophilic interactions. The closely related protein Tartan was partially redundant with Caps. These LRR proteins are probably part of a combinatorial cell-surface code that instructs discrete olfactory map formation. Nature Neuroscience 12, 1542 - 1550 (2009) Published online: 15 November 2009 http://www.nature.com/neuro/journal/v12/n12/abs/nn.2442.html?lang=en 59.Frequency Transitions in Odor-Evoked Neural Oscillations Iori Ito1, Maxim Bazhenov2, Rose Chik-ying Ong1, 3, Baranidharan Raman1, 4 and Mark Stopfer1, 1 National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, MD 20892, USA 34 2 Insect Frontiers, December 2009 Volume 1 Number 2 Department of Cell Biology and Neuroscience, University of California, Riverside, CA 92521, USA 3 Department of Biochemistry, The Chinese University of Hong Kong, Hong Kong 4 Chemical Science and Technology Laboratory, National Institute of Standards and Technology, 100 Bureau Drive, Stop 8362, Gaithersburg, MD 20899-8362, USA In many species, sensory stimuli elicit the oscillatory synchronization of groups of neurons. What determines the properties of these oscillations? In the olfactory system of the moth, we found that odors elicited oscillatory synchronization through a neural mechanism like that described in locust and Drosophila. During responses to long odor pulses, oscillations suddenly slowed as net olfactory receptor neuron (ORN) output decreased; thus, stimulus intensity appeared to determine oscillation frequency. However, changing the concentration of the odor had little effect upon oscillatory frequency. Our recordings in vivo and computational models based on these results suggested that the main effect of increasing odor concentration was to recruit additional, less well-tuned ORNs whose firing rates were tightly constrained by adaptation and saturation. Thus, in the periphery, concentration is encoded mainly by the size of the responsive ORN population, and oscillation frequency is set by the adaptation and saturation of this response. Neuron, Volume 64, Issue 5, 692-706, 10 December 2009 http://www.cell.com/neuron/abstract/S0896-6273%2809%2900805-8 60.Glycogen synthase kinase-3/Shaggy mediates ethanol-induced excitotoxic cell death of Drosophila olfactory neurons Rachael L. French1 and Ulrike Heberlein1 Department of Anatomy and Program in Neuroscience, University of California at San Francisco, 1550 4th Street, San Francisco, CA 94143-2324 It has long been known that heavy alcohol consumption leads to neuropathology and neuronal death. While the response of neurons to an ethanol insult is strongly influenced by genetic background, the underlying mechanisms are poorly understood. Here, we show that even a single intoxicating exposure to ethanol causes non-cell-autonomous apoptotic death specifically of Drosophila olfactory neurons, which is accompanied by a loss of a behavioral response to the smell of ethanol and a blackening of the third antennal segment. The Drosophila homolog of glycogen synthase kinase-3 (GSK-3)β, Shaggy, is required for ethanol-induced apoptosis. Consistent with this requirement, the GSK-3β inhibitor lithium protects against the neurotoxic effects of ethanol, indicating the possibility for pharmacological intervention in cases of alcohol-induced neurodegeneration. Ethanol-induced death of olfactory neurons requires both their neural activity and functional NMDA receptors. This system will allow the investigation of the genetic and molecular basis of ethanol-induced apoptosis in general and provide an understanding of the molecular role of GSK-3β in programmed cell death. PNAS December 8, 2009 vol. 106 no. 49 20924-20929 http://www.pnas.org/content/106/49/20924.abstract 61.Peripheral modulation of worker bee responses to queen mandibular pheromone Vanina Vergoz1, H. James McQuillan1, Lisa H. Geddes, Kiri Pullar, Brad J. Nicholson, Michael G. Paulin and Alison R. Mercer1 [email protected] Department of Zoology, University of Otago, P.O. Box 56, Dunedin 9054, New Zealand It is generally accepted that young worker bees (Apis mellifera L.) are highly attracted to queen mandibular pheromone (QMP). Our results challenge this widely held view. We have found that unless young workers are exposed to QMP early in adult life, they, like foragers, avoid contact with this pheromone. Our data indicate that responses to QMP are regulated peripherally, at the level of the antennal sensory neurons, and that a 35 Insect Frontiers, December 2009 Volume 1 Number 2 window of opportunity exists in which QMP can alter a young bee's response to this critically important pheromone. Exposing young bees to QMP from the time of adult emergence reduces expression in the antennae of the D1-like dopamine receptor gene, Amdop1. Levels of Amdop3 transcript, on the other hand, and of the octopamine receptor gene Amoa1, are significantly higher in the antennae of bees strongly attracted to QMP than in bees showing no attraction to this pheromone. A decline in QMP attraction with age is accompanied by a fall in expression in worker antennae of the D2-like dopamine receptor, AmDOP3, a receptor that is selectively activated by QMP. Taken together, our findings suggest that QMP's actions peripherally not only suppress avoidance behavior, but also enhance attraction to QMP, thereby facilitating attendance of the queen. PNAS December 8, 2009 vol. 106 no. 49 20930-20935 http://www.pnas.org/content/106/49/20930.abstract Top Insect Nutrition 62.Amino-acid imbalance explains extension of lifespan by dietary restriction in Drosophila Richard C. Grandison1,2, Matthew D. W. Piper1,2 & Linda Partridge1 [email protected] Institute of Healthy Ageing, Department of Genetics Evolution and Environment, University College London, Gower St, London WC1E 6BT, UK Dietary restriction extends healthy lifespan in diverse organisms and reduces fecundity1, 2. It is widely assumed to induce adaptive reallocation of nutrients from reproduction to somatic maintenance, aiding survival of food shortages in nature3, 4, 5, 6. If this were the case, long life under dietary restriction and high fecundity under full feeding would be mutually exclusive, through competition for the same limiting nutrients. Here we report a test of this idea in which we identified the nutrients producing the responses of lifespan and fecundity to dietary restriction in Drosophila. Adding essential amino acids to the dietary restriction condition increased fecundity and decreased lifespan, similar to the effects of full feeding, with other nutrients having little or no effect. However, methionine alone was necessary and sufficient to increase fecundity as much as did full feeding, but without reducing lifespan. Reallocation of nutrients therefore does not explain the responses to dietary restriction. Lifespan was decreased by the addition of amino acids, with an interaction between methionine and other essential amino acids having a key role. Hence, an imbalance in dietary amino acids away from the ratio optimal for reproduction shortens lifespan during full feeding and limits fecundity during dietary restriction. Reduced activity of the insulin/insulin-like growth factor signalling pathway extends lifespan in diverse organisms7, and we find that it also protects against the shortening of lifespan with full feeding. In other organisms, including mammals, it may be possible to obtain the benefits to lifespan of dietary restriction without incurring a reduction in fecundity, through a suitable balance of nutrients in the diet. Nature 462, 1061-1064 (24 December 2009) http://www.nature.com/nature/journal/v462/n7276/abs/nature08619.html?lang=en Top Insect Pharmacology 63.Transduction mechanism(s) of Na-saccharin in the blowfly Protophormia terraenovae: evidence for potassium and calcium conductance involvement Carla Masala1, Paolo Solari1, Giorgia Sollai1, Roberto Crnjar1 and Anna Liscia1 [email protected] 36 Insect Frontiers, December 2009 Volume 1 Number 2 1 Department of Experimental Biology, Section of General Physiology, University of Cagliari, Cittadella Universitaria di Monserrato, SS 554, Km 4.500, 09042 Monserrato, CA, Italy The study on transduction mechanisms underlying bitter stimuli is a particularly intriguing challenge for taste researchers. The present study investigates, in the labellar chemosensilla of the blowfly Protophormia terraenovae, the transduction mechanism by which saccharin evokes the response of the “deterrent” cell, with particular attention to the contribution of K+ and Ca2+ current and the role of cyclic nucleotides, since second messengers modulate Ca2+, Cl− and K+ currents to different extents. As assessed by extracellular single-sensillum recordings, our results show that the addition of a Ca2+ chelator such as EGTA or the Ca2+ current blockers SK&F-96365, Mibefradil, Nifedipine and W-7 decrease the response of the “deterrent” cell to saccharin. A similar decreasing effect was also obtained following the addition of 4-aminopyridine, a K+ current blocker. On the contrary, the membrane-permeable cyclic nucleotide 8-bromoguanosine 3′,5′cyclic monophosphate (8Br-cGMP) activates this cell and shows an additive effect when presented mixed with saccharin. Our results are consistent with the hypothesis that in the labellar chemosensilla of the blowfly both Ca2+ and K+ ions are involved in the transduction mechanism of the “deterrent” cell in response to saccharin. Our results also suggest a possible pathway common to saccharin and 8Br-cGMP. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1141-1151 http://www.springerlink.com/content/e423tu0uj7605u5u/ Top Insect Photoreception 64.Specialized ommatidia of the polarization-sensitive dorsal rim area in the eye of monarch butterflies have non-functional reflecting tapeta Thomas Labhart1 , Franziska Baumann1 and Gary D. [email protected] 1 Institute of Zoology, University of Zurich, Winterthurerstrasse 190, CH 8057 Zurich, Switzerland 2 Department of Electrical Engineering, University of Washington, Seattle, WA 98195-2500, USA Many insects exploit sky light polarization for navigation or cruising-course control. The detection of polarized sky light is mediated by the ommatidia of a small specialized part of the compound eye: the dorsal rim area (DRA). We describe the morphology and fine structure of the DRA in monarch butterflies (Danaus plexippus). The DRA consists of approximately 100 ommatidia forming a narrow ribbon along the dorsal eye margin. Each ommatidium contains two types of photoreceptor with mutually orthogonal microvilli orientations occurring in a 2:6 ratio. Within each rhabdomere, the microvilli are well aligned. Rhabdom structure and orientation remain constant at all retinal levels, but the rhabdom profiles, as seen in tangential sections through the DRA, change their orientations in a fan-like fashion from the frontal to the caudal end of the DRA. Whereas these properties (two microvillar orientations per rhabdom, microvillar alignment along rhabdomeres, ommatidial fan array) are typical for insect DRAs in general, we also report and discuss here a novel feature. The ommatidia of monarch butterflies are equipped with reflecting tapeta, which are directly connected to the proximal ends of the rhabdoms. Although tapeta are also present in the DRA, they are separated from the rhabdoms by a space of approximately 55 μm effectively inactivating them. This reduces self-screening effects, keeping polarization sensitivity of all photoreceptors of the DRA ommatidia both high and approximately equal. Cell and Tissue Research Volume 338, Number 3 / December, 2009 391-400 http://www.springerlink.com/content/h613q35p24540641/ 37 Insect Frontiers, December 2009 Volume 1 Number 2 Top Insect Physiology 65. Isoform- and cell-specific function of tyrosine decarboxylase in the Drosophila Malpighian tubule Edward M. Blumenthal [email protected] Department of Biological Sciences, Marquette University, P.O. Box 1881, Milwaukee, WI 53201-1881, USA The biogenic amine tyramine (TA) is a potent diuretic factor when applied to the Malpighian tubule (MT) of Drosophila melanogaster, stimulating both urine production and transepithelial chloride conductance. Isolated MTs can respond not only to TA but also to its precursor, tyrosine; this observation led to the proposal that MTs are able to synthesize TA from applied tyrosine through the action of the enzyme tyrosine decarboxylase (TDC). In the current study it is shown that the non-neuronal isoform of TDC, Tdc1, is expressed in the principal cells of the MT. A mutant allele of Tdc1, Tdc1f03311, was identified that reduced expression of the mature Tdc1 transcript by greater than 100-fold. MTs isolated from Tdc1f03311 homozygous flies showed no significant depolarization of their transepithelial potential (TEP) or diuresis in response to tyrosine while retaining normal sensitivity to TA. By contrast, a previously identified null mutant allele of the neuronal TDC isoform Tdc2 had no effect on either tyrosine or TA sensitivity. To determine in which cell type of the MT Tdc1 expression is required, flies were generated carrying a UAS-Tdc1 transgene and cell-type-specific Gal4 drivers on a Tdc1f03311 homozygous background. Rescue of Tdc1 expression in principal cells fully restored sensitivity to tyrosine whereas expression of Tdc1 in stellate cells had no rescuing effect. It is concluded that synthesis of TA by Tdc1 in the principal cells of the MT is required for physiological responses to tyrosine. TA synthesis in the MT is the first reported physiological role for Drosophila Tdc1. Journal of Experimental Biology 212, 3802-3809 (2009) http://jeb.biologists.org/cgi/content/abstract/212/23/3802?etoc Top Insect Sex Determination 66.The origin of a selfish B chromosome triggering paternal sex ratio in the parasitoid wasp Trichogramma kaykai Joke J. F. A. Van Vugt1,†Hans de Jong2 and Richard Stouthamer3,* [email protected] 1. Laboratory of Entomology, Wageningen University, PO Box 8031, 6700 EH Wageningen, The Netherlands 2. Laboratory of Genetics, Wageningen University, Arboretumlaan 4, 6703 BD Wageningen, The Netherlands 3. Department of Entomology, University of California, Riverside, CA 92521, USA This study uses molecular and cytogenetic methods to determine the origin of a B chromosome in some males of the wasp Trichogramma kaykai. This so-called paternal sex ratio (PSR) chromosome transmits only through sperm and shortly after fertilization triggers degeneration of the paternal genome, while keeping itself intact. The resulting embryos develop into haploid B-chromosome-carrying males. Another PSR chromosome with a very similar mode of action is found in the distantly related wasp Nasonia vitripennis and its origin was traced by transposon similarity to the genus Trichomalopsis, which is closely related to Nasonia. To determine whether both PSR chromosomes have a similar origin we aimed to reveal the origin of the Trichogramma PSR chromosome. Using fluorescent in situ hybridization, we discovered a major satellite repeat on the PSR chromosome, the 45S ribosomal DNA. Analysis of the internal transcribed spacer 2 (ITS2) of this repeat showed the presence of multiple ITS2 sequences on the PSR chromosome 38 Insect Frontiers, December 2009 Volume 1 Number 2 resembling either the ITS2 of T. oleae or of T. kaykai. We therefore conclude that the Trichogramma PSR chromosome originates from T. oleae or a T. oleae-like species. Our results are consistent with different origins for the PSR chromosomes in Trichogramma and Nasonia. Proc. R. Soc. B 7 December 2009 vol. 276 no. 1676 4149-4154 http://rspb.royalsocietypublishing.org/content/276/1676/4149.abstract Top Insect Society 67. Brood pheromone suppresses physiology of extreme longevity in honeybees (Apis mellifera) B. Smedal1, M. Brynem2, C. D. Kreibich1 and G. V. Amdam1,3,* [email protected] 1 Department of Chemistry, Biotechnology and Food Science, University of Life Sciences, P.O. Box 5003, N-1432 Aas, Norway 2 Department of Animal and Aquacultural Sciences, University of Life Sciences, P.O. Box 5003, N-1432 Aas, Norway 3 School of Life Science, Arizona State University, Tempe, P.O. Box 874501, AZ 85287, USA. Honeybee (Apis mellifera) society is characterized by a helper caste of essentially sterile female bees called workers. Workers show striking changes in lifespan that correlate with changes in colony demography. When rearing sibling sisters (brood), workers survive for 3-6 weeks. When brood rearing declines, worker lifespan is 20 weeks or longer. Insects can survive unfavorable periods on endogenous stores of protein and lipid. The glycolipoprotein vitellogenin extends worker bee lifespan by functioning in free radical defense, immunity and behavioral control. Workers use vitellogenin in brood food synthesis, and the metabolic cost of brood rearing (nurse load) may consume vitellogenin stores and reduce worker longevity. Yet, in addition to consuming resources, brood secretes a primer pheromone that affects worker physiology and behavior. Odors and odor perception can influence invertebrate longevity but it is unknown whether brood pheromone modulates vitellogenin stores and survival. We address this question with a 2factorial experiment where 12 colonies are exposed to combinations of absence vs presence of brood and brood pheromone. Over an age-course of 24 days, we monitor the amount of vitellogenin stored in workers' fat body (adipose tissue). Thereafter, we track colony survival for 200 days. We demonstrate that brood rearing reduces worker vitellogenin stores and colony long-term survival. Yet also, we establish that the effects can result solely from exposure to brood pheromone. These findings indicate that molecular systems of extreme lifespan regulation are integrated with the sensory system of honeybees to respond to variation in a primer pheromone secreted from larvae. Journal of Experimental Biology 212, 3795-3801 (2009) http://jeb.biologists.org/cgi/content/abstract/212/23/3795?etoc 68.Flexible task allocation and the organization of work in ants Elva J. H. Robinson1,*Ofer Feinerman2 and Nigel R. Franks1 [email protected] School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, UK Computational Biology Center, Memorial Sloan-Kettering Cancer Center, 1275 York Avenue, Box 460, New York, NY 10021, USA Flexibility in task performance is essential for a robust system of division of labour. We investigated what factors determine which social insect workers respond to colony-level changes in task demand. We used radio-frequency identification technology to compare the roles of corpulence, age, spatial location and previous activity (intra-nest/extra-nest) in determining whether worker ants (Temnothorax albipennis) respond to an increase in demand for foraging or brood care. The less corpulent ants took on the extra foraging, 39 Insect Frontiers, December 2009 Volume 1 Number 2 irrespective of their age, previous activity or location in the nest, supporting a physiological threshold model. We found no relationship between ants that tended the extra brood and corpulence, age, spatial location or previous activity, but ants that transported the extra brood to the main brood pile were less corpulent and had high previous intra-nest activity. This supports spatial task-encounter and physiological threshold models for brood transport. Our data suggest a flexible task-allocation system allowing the colony to respond rapidly to changing needs, using a simple task-encounter system for generalized tasks, combined with physiologically based response thresholds for more specialized tasks. This could provide a social insect colony with a robust division of labour, flexibly allocating the workforce in response to current needs. Proc. R. Soc. B 22 December 2009 vol. 276 no. 1677 4373-4380 http://rspb.royalsocietypublishing.org/content/276/1677/4373.abstract 69.Polymorphic social organization in an ant Richard J. Gill1, Andres Arce1, Laurent Keller2 and Robert L. Hammond1,* [email protected] 1. Department of Biological Sciences, University of Hull, Cottingham Road, Hull HU6 7RX, UK 2. Department of Ecology and Evolution, Biophore, University of Lausanne, 1015 Lausanne, Switzerland Identifying species exhibiting variation in social organization is an important step towards explaining the genetic and environmental factors underlying social evolution. In most studied populations of the ant Leptothorax acervorum, reproduction is shared among queens in multiple queen colonies (polygyny). By contrast, reports from other populations, but based on weaker evidence, suggest a single queen may monopolize all reproduction in multiple queen colonies (functional monogyny). Here we identify a marked polymorphism in social organization in this species, by conclusively showing that functional monogyny is exhibited in a Spanish population, showing that the social organization is stable and not purely a consequence of daughter queens overwintering, that daughter queen re-adoption is frequent and queen turnover is low. Importantly, we show that polygynous and functionally monogynous populations are not genetically distinct from one another based on mtDNA and nDNA. This suggests a recent evolutionary divergence between social phenotypes. Finally, when functionally monogynous and polygynous colonies were kept under identical laboratory conditions, social organization did not change, suggesting a genetic basis for the polymorphism. We discuss the implications of these findings to the study of reproductive skew. Proc. R. Soc. B 22 December 2009 vol. 276 no. 1677 4423-4431 http://rspb.royalsocietypublishing.org/content/276/1677/4423.abstract Top Insect Vision 70.Visual detection of diminutive floral guides in the bumblebee Bombus terrestris and in the honeybee Apis mellifera Klaus Lunau1 , Katrin Unseld1 and Franziska Wolter1 [email protected] 1 Sensory Ecology Group, Institute of Neurobiology, Heinrich-Heine-University, 40225 Düsseldorf, Germany Many flowers display colour patterns comprising a large peripheral colour area that serves to attract flower visitors from some distance, and a small central, contrastingly coloured area made up by stamens or floral guides. In this study, we scaled down the size of floral guides to detect the minimal size bumblebees (Bombus terrestris) and honeybees (Apis mellifera) require for guidance. We analyzed the approach and the precise contact of the antennal tips with the floral guide of artificial flowers which precedes landing and inspection. Both bumblebees and honeybees were able to make 40 Insect Frontiers, December 2009 Volume 1 Number 2 antennal contact with circular floral guides which were 2 mm in diameter; bumblebees performed better than honeybees and antennated also at floral guides smaller than 2 mm. In discrimination experiments with bumblebees, a minimum floral guide size of 2 mm was required for discrimination between artificial flowers with and without floral guides. With increasing experience bumblebees targeted close to the site of reward instead of making antennal contact with the floral guide, whereas honeybees did not alter their initial behaviour with growing experience. Bumblebees and honeybees spontaneously target diminutive floral guides to achieve physical contact with flowers by means of their antennae which helps them to inspect flowers. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology Volume 195, Number 12 / December, 2009 1121-1130 http://www.springerlink.com/content/tg2g7127654hjxn6/ Top Insect Review 71.Are Bigger Brains Better? Lars Chittka1, and Jeremy Niven2 1 Queen Mary University of London, Research Centre for Psychology, School of Biological and Chemical Sciences, Mile End Road, London E1 4NS, UK 2 University of Cambridge, Department of Zoology, Downing Street, Cambridge CB2 3EJ, UK Attempts to relate brain size to behaviour and cognition have rarely integrated information from insects with that from vertebrates. Many insects, however, demonstrate that highly differentiated motor repertoires, extensive social structures and cognition are possible with very small brains, emphasising that we need to understand the neural circuits, not just the size of brain regions, which underlie these feats. Neural network analyses show that cognitive features found in insects, such as numerosity, attention and categorisation-like processes, may require only very limited neuron numbers. Thus, brain size may have less of a relationship with behavioural repertoire and cognitive capacity than generally assumed, prompting the question of what large brains are for. Larger brains are, at least partly, a consequence of larger neurons that are necessary in large animals due to basic biophysical constraints. They also contain greater replication of neuronal circuits, adding precision to sensory processes, detail to perception, more parallel processing and enlarged storage capacity. Yet, these advantages are unlikely to produce the qualitative shifts in behaviour that are often assumed to accompany increased brain size. Instead, modularity and interconnectivity may be more important Current Biology, Volume 19, Issue 21, R995-R1008, 17 November 2009 http://www.cell.com/current-biology/abstract/S0960-9822(09)01597-8 72.A Review of Ant Cuticular Hydrocarbons Stephen Martin1 and Falko Drijfhout2 [email protected] 1. Department of Animal and Plant Sciences, University of Sheffield, Sheffield, UK 2. Chemical Ecology Group, School of Physical and Geographical Sciences, Lennard-Jones Laboratory, Keele University, Keele, UK We compared the published cuticular hydrocarbon (CHC) profiles of 78 ant species across 5 subfamilies. Almost 1,000 CHCs have been described for these species, composing 187 distinct homologous series and ten hydrocarbon groups. In descending order of occurrence were: n-alkanes > monomethylalkanes > dimethylalkanes > alkenes > dienes>> trimethylalkanes>> methylalkenes > methylalkadienes > trienes > tetramethylalkanes. Odd chain lengths and positions of methyl or double bonds at odd carbon numbers were far more numerous than even chain-length compounds or bond positions. Although each species possess its own unique pattern of CHCs, we found no 41 Insect Frontiers, December 2009 Volume 1 Number 2 association between CHC profile and phylogeny. The production of the biosynthetically complex compounds (e.g., methyl branched dienes) by the most primitive living ant suggests that the basic genetic architecture required to produce the rich diversity of CHCs was already present prior to their adaptive radiation. Unlike the ubiquitous nalkanes and monomethylalkanes, there is a huge diversity of species-specific dimethylalkanes that makes them likely candidates for species and nest-mate discrimination signals. Journal of Chemical Ecology Volume 35, Number 10 / October, 2009 1151-1161 http://www.springerlink.com/content/h475723350505151/ 73.Courtship learning in Drosophila melanogaster: Diverse plasticity of a reproductive behavior Leslie C. Griffith1 and Aki Ejima2 [email protected] Department of Biology, National Center for Behavioral Genomics and Volen Center for Complex Systems, Brandeis University, Waltham, Massachusetts 02493, USA; Career-Path Promotion Unit for Young Life Scientists, Kyoto University, Kyoto, Japan 606-8501 Mechanisms for identifying appropriate mating partners are critical for species propagation. In many species, the male uses multiple sensory modalities to search for females and to subsequently determine if they are fit and receptive. Males can also use the information they acquire in this process to change their courtship behavior and reduce courtship of classes of targets that are inappropriate or unreceptive. In Drosophila, courtship plasticity, in the form of both nonassociative and associative learning, has been documented—the type of learning depending on the nature of the trainer. The conditions in which the male is presented with the training target can profoundly alter the cues that he finds salient and the longevity of the memory that he forms. With the exception of habituation and sensitization, these types of plasticity have an operant component in that the male must be courting to respond to the behavior-altering cues. Courtship plasticity is therefore a complex and rich range of behaviors rather than a single entity. Our understanding of these plastic behaviors has been enhanced by recent advances in our understanding of the circuitry underlying courtship itself and the identification of chemical cues that drive and modify the behavior. Courtship learning is providing a window into how animals can use a variety of sensory inputs to modulate a decision making process at many levels. Learn. Mem. 2009. 16: 743-750 http://learnmem.cshlp.org/content/16/12/743.abstract 74.Circadian rhythms and the evolution of photoperiodic timing in insects DAVID S. SAUNDERS Emeritus Professor, University of Edinburgh, Scotland, U.K. This review discusses possible evolutionary trends in insect photoperiodism, mainly from a chronobiological perspective. A crucial step was the forging of a link between the hormones regulating diapause and the systems of biological rhythms, circadian or circannual, which have independently evolved in eukaryotes to synchronize physiology and behaviour to the daily cycles of light and darkness. In many of these responses a central feature is that the circadian system resets to a constant phase at the beginning of the subjective night, and then 'measures' the duration of the next scotophase. In 'external coincidence', one version of such a clock, light now has a dual role. First, it serves to entrain the circadian system to the stream of pulses making up the light/dark cycle and, second, it regulates the nondiapause/diapause switch in development by illuminating/not illuminating a specific light sensitive phase falling at the end of the critical night length. Important work by A. D. Lees on the aphid Megoura viciae using socalled 'night interruption experiments' demonstrates that pulses falling early in the night 42 Insect Frontiers, December 2009 Volume 1 Number 2 lead to long-day effects that are reversible by a subsequent dark period longer than the critical night length and also show maximal sensitivity in the blue–green range of the spectrum. Pulses falling in the latter half of the night, however, produce long-day effects that are irreversible by a subsequent long-night and show a spectral sensitivity extending into the red. With movement to higher latitudes, insects develop genetic clines in various parameters, including critical night length, the number of long-night cycles needed for diapause induction, the strength of the response, and the 'depth' or intensity of the diapause thus induced. Evidence for these and other types of photoperiodic response suggests that they provided strong selective advantages for insect survival. Physiological Entomology Volume 34 Issue 4, Pages 301 – 308 Published Online: 19 Nov 2009 http://www3.interscience.wiley.com/journal/122688579/abstract Top Edited by Xin-Cheng Zhao 2009-12-30 43