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Acknowledgements We thank Michael Steinert and Silke Killinger of the Institute for Molecular Infection Biology of the University of Würzburg for help and introduction to the FISH technique. We would like to thank Roy Gross for helpful suggestions and discussions, and Norbert Weißel for comments on the manuscript. We thank Jon Seger for providing us with P. gibbosus specimens, Johannes Kroiss for collecting P. venustus in Greece, Thomas Eltz for sending P. triangulum from Düsseldorf, Silke Fregin for collecting P. triangulum in Schweinfurt, and Protsenko Yura for sending P. triangulum specimens from the Ukraine. 2.4 REFERENCES Amann, R.I., Binder, B.J., Olson, R.J., Chisholm, S.W., Devereux, R. & Stahl, D.A. 1990. Combination of 16S ribosomal RNA targeted oligonucleotide probes with flow-cytometry for analyzing mixed microbial populations. Appl. Environ. Microbiol. 56 (6): 1919-1925. Behal, V. 2000. Bioactive products from Streptomyces. In. Advances in Applied Microbiology: 113-156. New York: Academic Press. Buchner, P. 1921. Tier und Pflanze in intrazellulärer Symbiose. Berlin: Verlag Gebrüder Borntraeger. Currie, C.R. 2001. A community of ants, fungi, and bacteria: A multilateral approach to studying symbiosis. Annu. Rev. Microbiol. 55: 357-380. Currie, C.R., Bot, A.N.M. & Boomsma, J.J. 2003. Experimental evidence of a tripartite mutualism: bacteria protect ant fungus gardens from specialized parasites. Oikos 101 (1): 91-102. Currie, C.R., Scott, J.A., Summerbell, R.C. & Malloch, D. 1999. Fungus-growing ants use antibioticproducing bacteria to control garden parasites. Nature 398 (6729): 701-704. Goodfellow, M. & Cross, T. 1984. The biology of actinomycetes. London: Academic. Grimm, D., Merkert, H., Ludwig, W., Schleifer, K.H., Hacker, J. & Brand, B.C. 1998. Specific detection of Legionella pneumophila: Construction of a new 16S rRNA-targeted oligonucleotide probe. Appl. Environ. Microbiol. 64: 2686-2690. Kutzner, H.J. 1981. The family Streptomycetaceae. In: Starr, M.P., Stolp, H., Trüper, H.G., Balows, A. & Schlegel, H.G. (eds.). The Prokaryotes: 2028-2090. Berlin: Springer. Maidak, B.L., Cole, J.R., Lilburn, T.G., Parker, C.T., Saxman, P.R., Farris, R.J., Garrity, G.M., Olsen, G.J., Schmidt, T.M. & Tiedje, J.M. 2001. The RDP-II (Ribosomal Database Project). Nucleic Acids Res. 29 (1): 173-174. Margulis, L. & Fester, R. 1991. Symbiosis as a source of evolutionary innovation: speciation and morphogenesis. Cambridge: MIT Press. Maynard-Smith, J. 1989. Generating novelty by symbiosis. Nature 341: 284-285. Moran, N.A. & Baumann, P. 2000. Bacterial endosymbionts in animals. Curr. Opin. Microbiol. 3: 270- 275. Pernodet, J.L., Boccard, F., Alegre, M.T., Gagnat, J. & Guerineau, M. 1989. Organization and nucleotide sequence analysis of a ribosomal RNA gene cluster from Streptomyces ambofaciens. Gene 79 (1): 33-46. Poulsen, M., Bot, A.N.M., Currie, C.R., Nielsen, M.G. & Boomsma, J.J. 2003. Within-colony transmission and the cost of a mutualistic bacterium in the leaf-cutting ant Acromyrmex octospinosus. Funct. Ecol. 17 (2): 260-269. Rintala, H., Nevalainen, A., Ronka, E. & Suutari, M. 2001. PCR primers targeting the 16S rRNA gene for the specific detection of streptomycetes. Mol. Cell. Probes 15 (6): 337-347. Sapp, J. 1994. Evolution by association: a history of symbiosis. New York: Oxford University Press. Sauer, C., Dudaczek, D., Holldobler, B. & Gross, R. 2002. Tissue localization of the endosymbiotic bacterium 'Candidatus Blochmannia floridanus' in adults and larvae of the carpenter ant Camponotus floridanus. Appl. Environ. Microbiol. 68 (9): 4187-4193. Stach, J.E.M., Maldonado, L.A., Ward, A.C., Goodfellow, M. & Bull, A.T. 2003. New primers for the class Actinobacteria: application to marine and terrestrial environments. Env. Microbiol. 5 (10): 828-841. 44
Strohm, E. 1995. Allokation elterlicher Investitionen beim Europäischen Bienenwolf Philanthus triangulum Fabricius (Hymenoptera: Sphecidae). Berlin: Verlag Dr. Köster. Strohm, E. & Linsenmair, K.E. 1995. Leaving the cradle - how beewolves (Philanthus triangulum F.) obtain the necessary spatial information for emergence. Zoology 98 (3): 137-146. Strohm, E. & Linsenmair, K.E. 1999. Measurement of parental investment and sex allocation in the European beewolf Philanthus triangulum F. (Hymenoptera: Sphecidae). Behav. Ecol. Sociobiol. 47 (1-2): 76-88. Weisburg, W.G., Barns, S.M., Pelletier, D.A. & Lane, D.J. 1991. 16s ribosomal DNA amplification for phylogenetic study. J. Bacteriol. 173 (2): 697-703. Accession numbers Partial 16S rDNA sequences from Streptomyces endosymbionts of Philanthus triangulum (from three different populations: Würzburg, Germany; Schweinfurt, Germany; and from the Ukraine), P. venustus and P. gibbosus are available at GenBank (http://www.ncbi.nlm.nih.gov/) with the accession numbers AY854952-AY854956. 45
Page 2 and 3: Protective bacteria and attractive
Page 5: “We are symbionts on a symbiotic
Page 8 and 9: CHAPTER 4: MORPHOLOGY AND ULTRASTRU
Page 10 and 11: 8.4 Results........................
Page 13: LIST OF PUBLICATIONS This thesis is
Page 16 and 17: major factor driving speciation and
Page 18 and 19: et al. 2002; Gil et al. 2003; Rio e
Page 20 and 21: eetles use for protection against p
Page 22 and 23: either benefit directly, if males v
Page 24 and 25: (Arnold 1925; Erlandsson 1962; Boha
Page 26 and 27: changing the direction of their bod
Page 28 and 29: Therefore, “bad” genes from the
Page 30 and 31: allow the assessment of male reprod
Page 32 and 33: Currie, C.R., Mueller, U.G. & Mallo
Page 34 and 35: Herzner, G., Schmitt, T., Linsenmai
Page 36 and 37: Mueller, U.G., Rehner, S.A. & Schul
Page 38 and 39: Stouthamer, R., Breeuwer, J.A.J. &
Page 41 and 42: CHAPTER 2 SYMBIOTIC BACTERIA PROTEC
Page 43 and 44: A promising candidate for such a pr
Page 45 and 46: actinomycete bacteria of the family
Page 47 and 48: unlikely that this association is l
Page 49: unspecific eubacterial probe EUB 33
Page 54 and 55: 3.3 ERGEBNISSE UND DISKUSSION Der B
Page 56 and 57: Aus evolutionären und ökologische
Page 59 and 60: CHAPTER 4 MORPHOLOGY AND ULTRASTRUC
Page 61 and 62: gland secretions are applied onto f
Page 63 and 64: gland cell units (according to Noir
Page 65 and 66: (fl) covering the reservoir opening
Page 67 and 68: 4.4.2 Ultrastructure Scanning elect
Page 69 and 70: (C) Plasma membrane invaginations (
Page 71 and 72: e sequestered from the hemolymph, a
Page 73: Herzner, G., Schmitt, T., Linsenmai
Page 76 and 77: 5.2 INTRODUCTION Many insects have
Page 78 and 79: 5.3.3 DNA extraction, PCR and seque
Page 80 and 81: glands from which they are secreted
Page 82 and 83: The bacteria were clearly stained b
Page 84 and 85: actinomycete primers (Act-S20 and A
Page 86 and 87: endosymbionts are needed to confirm
Page 88 and 89: µg/ml), and beewolf cocoon agar (a
Page 90 and 91: CaCl2 MgCl2 0.15 g 0.05 g KCl 0.2 g
Page 92 and 93: Nyholm, S.V., Stabb, E.V., Ruby, E.
Page 94 and 95: 6.2 INTRODUCTION Many insects harbo
Page 96 and 97: 6.3.2 Gas chromatography - mass spe
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Discriminant function 2 2 1 0 -1 -2
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6.5 DISCUSSION GC-MS analyses of th
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6.6 REFERENCES Aitchison, J. 1986.
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100
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7.2 INTRODUCTION Hymenoptera posses
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Cuticular hydrocarbons were obtaine
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Table 7.1: List of compounds in the
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(notably pentacosene and heptacosen
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unicellular epithelium (Strohm et a
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Dahbi, A., Cerda, X. & Lenoir, A. 1
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114
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8.2 INTRODUCTION Senescence is a pr
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heavily depends on her “quality
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transformed absolute amounts of the
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(S)-2,3-Dihydrofarnesoic acid acid
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Dihydrofarnesoic acid, nonadecenone
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Jones, T.M. & Hamilton, J.G.C. 1998
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9.2 INTRODUCTION Male sexual signal
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al. 2003; Kroiss et al. submitted).
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and kept in small polystyrol vials
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heptacosene > 0.55) (Fig. 9.1) (Kro
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pentacosen-8-one (C25one); a mixtur
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Populations in both data sets and,
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Table 9.2: Relative effects of fami
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9.5 DISCUSSION In the present study
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If female beewolves use the informa
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Herzner, G., Goettler, W., Kroiss,
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Strohm, E., Laurien-Kehnen, C. & Bo
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within their territory that is like
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to 0.900 (Table 10.1). Thus, there
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154
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Bin 1995; Bin et al. 1999; Isidoro
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symbionts are transmitted verticall
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vitamins and in sexual maturation o
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Additionally, the genome sequence o
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Optimal outbreeding The degree of k
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investigate the importance of in- a
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Age has an effect on both the amoun
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to confirm these results, a connect
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Baumann, L. & Baumann, P. 2005. Cos
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endosymbiont Streptomyces anulatus
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Lopez, P., Aragon, P. & Martin, J.
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Sagvik, J., Uller, T. & Olsson, M.
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180
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could be detected in closely relate
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184
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Genetische Analysen zeigen, dass di
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188
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Der Arbeitsgruppe Gadau/Feldhaar da
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192
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WS2005/06 Fortgeschrittenen-Praktik
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