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Sexual Cooperation in Two Male-Diphenic Cardiocondyla Species

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Sexual Cooperation in Two Male-Diphenic Cardiocondyla Species Sexual cooperation in two male-diphenic Cardiocondyla species DISSERTATION ZUR ERLANGUNG DES DOKTORGRADES DER NATURWISSENSCHAFTEN (DR. RER. NAT.) DER FAKULTÄT FÜR BIOLOGIE UND VORKLINISCHE MEDIZIN DER UNIVERSITÄT REGENSBURG vorgelegt von Marion Füßl aus Ingolstadt im Jahr 2014 Sexual cooperation in two male-diphenic Cardiocondyla species DISSERTATION ZUR ERLANGUNG DES DOKTORGRADES DER NATURWISSENSCHAFTEN (DR. RER. NAT.) DER FAKULTÄT FÜR BIOLOGIE UND VORKLINISCHE MEDIZIN DER UNIVERSITÄT REGENSBURG vorgelegt von Marion Füßl aus Ingolstadt im Jahr 2014 Promotionsgesuch eingereicht am: 19.12.2014 Die Arbeit wurde angeleitet von Dr. A. Schrempf Prüfungsausschuss: Vorsitzender: Prof. Dr. S. Schneuwly 1. Gutachter: Prof. Dr. J. Heinze 2. Gutachter: Prof. Dr. J. Ruther 3. Prüfer: Prof. Dr. C. Oberprieler Unterschrift: If you watch animals objectively for any length of time, you’re driven to the conclusion that their main aim in life is to pass on their genes to the next generation. – David Attenborough – Manuscripts that contributed to this Thesis and Author’s contribution Fuessl M, Santos CG, Bayersdorfer F, Skorupa J, Hartfelder KH, Heinze J, Schrempf A (in preparation) Suppression subtractive hybridization reveals differentially expressed genes in the accessory glands of two ant male phenes with different life histories, (Chapter 2) Author’s contribution: JH, KHH and AS designed and supervised the study. FB gave helpful advice for qPCR. AS contributed to the manuscript. MF and CGS performed the RDA. JS performed the qPCR which was supervised by MF and AS. Bioinformatic analysis to obtain USs were done by CGS and database searches by MF. Dissection, cDNA-synthesis, primer design, sample preparations, evaluation of qPCR results and writing the draft of the manuscript was done by MF. Fuessl M, Reinders J, Oefner PJ, Heinze J, Schrempf A (in preparation) Comparison of the accessory gland proteins of two male-diphenic ant species and the effect on their mating partners. (Chapter 3) Author’s contribution: JH and AS supervised the study. PJO, JR and AS contributed to the manuscript. JR performed the nano-LC-MS/MS-analysis and helped with evaluation of results. Dissection, sample preparation, IEF, 2D-SDS-PAGE, silver-staining and writing the draft of the manuscript was done by MF. Fuessl M, Reinders J , Oefner PJ, Heinze J, Schrempf A (2014) Selenophosphate synthetase in the male accessory glands of an insect without selenoproteins. Journal of Insect Physiology. 71: 46-51 (Chapter 4) Author’s contribution: JH and AS supervised the study. PJO, JR and AS gave helpful comments on the manuscript. JR performed the nano-LC-MS/MS-analysis and helped with evaluation of results. Dissection, sample preparation, IEF, 2D-SDS-PAGE, silver- staining and writing the draft of the manuscrupt was done by MF. Fuessl M, Heinze J, Schrempf A (accepted for publication) Queen and male longevity in the Southeast Asian ant Cardiocondyla tjibodana Karavaiev, 1935. Journal of Asian Myrmecology. (Chapter 5) Author’s contribution: AS designed and supervised the study. Data was collected by MF. AS, JH and MF evaluated the data. MF wrote the first draft of the manuscript, on which JH developed the final manuscript. ii Contents Manuscripts that contributed to this Thesis and Author’s contribution ii Chapter 1: General Introduction 1 1.1 Early theories of the evolution of eusociality . .1 1.2 Eusocial insect life history . .2 1.3 Seminal fluid, accessory gland proteins and longevity . .3 1.4 The study species Cardiocondyla obscurior and Cardiocondyla tjibodana: their life-cycle and its male phenes . .6 1.5 Ageing theories, a very brief overview . 10 1.6 Aims of the Thesis . 12 Chapter 2: Suppression subtractive hybridization reveals differentially expressed genes in the accessory glands of two ant male phenes with different life histories* 15 2.1 Introduction . 17 2.2 Material and Methods . 20 2.2.1 Colony housing and male dissection . 20 2.2.2 Representational difference analysis . 21 2.2.3 Bioinformatic analysis . 22 2.2.4 Differential gene expression analysis by quantitative RT-PCR . 22 2.3 Results . 23 2.3.1 RDA library characteristics and bioinformatic analyses . 23 2.3.2 Quantitative RT-PCR analysis of RDA-library genes . 24 2.4 Discussion . 29 Chapter 3: Comparison of the accessory gland proteins of two male- diphenic ant species and the effect on their mating partners* 35 3.1 Introduction . 37 3.2 Material and Methods . 40 3.2.1 Ant sampling and housing . 40 3.2.2 Preparation of samples and performance of the first dimension: Isoelectric focusing (IEF) . 40 3.2.3 Sample equilibration and performance of the second dimension (SDS-PAGE) . 41 3.2.4 Silver staining . 42 3.2.5 Nano-LC-MS/MS-analysis . 42 3.3 Results . 42 3.4 Discussion . 50 iii Contents Chapter 4: Selenophosphate synthetase in the male accessory glands of an insect without selenoproteins 55 4.1 Introduction . 57 4.2 Material and methods . 58 4.2.1 Ant sampling and housing . 58 4.2.2 Sample preparation and performance of the first dimension: Iso- electric focusing (IEF) . 58 4.2.3 Preparation and performance of the second dimension: SDS–PAGE 59 4.2.4 Silver staining . 60 4.2.5 Nano-LC–MS/MS-analysis . 60 4.2.6 PCR . 60 4.3 Results . 60 4.4 Discussion . 63 Chapter 5: Queen and male longevity in the Southeast Asian ant Car- diocondyla tjibodana Karavaiev, 1935 67 5.1 Introduction . 69 5.2 Material and Methods . 69 5.3 Results and Discussion . 70 Chapter 6: General Discussion 75 6.1 From the RNA to the protein: a brief overview . 76 6.2 SERPINB10 a potential factor in the insect immune system (Chapters 2 & 3) 77 6.3 The ejaculatory bulb-specific protein 3-like as interesting result of the RDA (Chapter 2) . 78 6.4 The evolution of SELD/SPS1 and its possible relation to selenoprotein biosynthesis, ROS & ageing (Chapters 3 & 4) . 78 6.5 PDIA3 a thiol oxidoreductase (Chapter 3) . 81 6.6 Changing composition of the seminal fluid and how the ACPs might act in C. obscurior ................................... 81 6.7 How the ACPs of C. tjibodana might be influenced by the mating system (Chapters 3 & 5) . 83 6.8 The ageing theories with special emphasis on social insects . 84 Chapter 7: Summary 87 Chapter 8: Zusammenfassung 89 Danksagung 91 Bibliography 93 Eidesstattliche Erklärung 128 iv Abbreviations (except units, constants and chemical elements) Name of a species: first letter of the genus in capital letter plus name of the species in lower-case, both in italics. Example: Cardiocondyla (genus) obscurior (species) = C. obscurior DROME Drosophila melanogaster (exclusively Chapter 2, otherwise D. melanogaster) CAROB Cardiocondyla obscurior (exclusively Chapter 2, otherwise C. obscurior) APIME Apis mellifera (exclusively Chapter 2, otherwise A. mellifera) 2D-SDS-PAGE two-dimensional-sodiumdodecylsulphate-polyacrylmide gel electrophoresis ACP accessory gland protein ANOVA analysis of variance ATP adenosine triphosphate blast basic logical alignement search tool bp basepair cDNA complemetary DNA CHAPS 3-[(3-Cholamidopropyl) dimethylammonio]-1-propanesulfonate Cq quantification cycle (= treashold cycle Ct) CRISP cysteine rich secretory proteins DNA deoxyribonucleic acid DTT DL dithiothreitol EBSP see PEB-me ER endoplasmatic reticulum v Abbreviations EST expressed sequence tag fcv fold change value Ghitm Growth-hormone inducible transmembrane protein IAA iodoacetamide IEF isoelectric focusing IPG immobilized pH gradient IPTG isoproyl β-D-1-thiogalactopyranoside LB lysogeny broth LC-MS/MS-analysis liquid chromatography-mass spectrometry/mass spectrometry- analysis LRR Leucin rich repeat MAG male accessory gland mRNA messenger RNA NCBI national center of biotechnology information NHERF Na+/H+ exchange regulatory cofactor NL non-linear nr non-redundant PBS protein biosynthesis PCR polymerase chain reaction PDIA3 protein disulfide isomerase A3 PEB-me ejaculatory bulb specific protein (of Drosophila melanogaster) PSA prostate specific antigen ptuf see SPS qPCR quantitative real-time PCR RDA representational difference analysis RNA ribonucleic acid vi Abbreviations ROS reactive oxygen species SECIS selenocysteine insertion sequence SELA-D cofactors in bacterial selenoproteine-biosynthesis SERPIN serine protease inhibitor SFP seminal fluid protein SOD superoxide dismutase SP sex peptide SPS, sps, ptuf selenophosphate synthetase SRY sex-determining region Y Tris-HCl tris(hydroxymethyl)-aminomethane US unique sequence UTR untranslated region v/v volume per volume w/v weight per volume vii Chapter 1 General Introduction 1.1 Early theories of the evolution of eusociality When Darwin first published his book "on the origin of species by means of natural selection" in the year 1859 [Darwin, 1859], he wouldn’t obscure that his theory of natural selection still posed him an enormous riddle: the sterile female worker caste of social insects. Why would an individual forgo its own reproduction, or even die for the sake of the colony? And how could these workers be so different from fertile males and females concerning "morphology and instinct"? He was puzzled by "how such correlated modifications of structure could have been slowly accumulated by natural selection", as sterile workers "could never have transmitted successively acquired modifications of structure or instinct to its progeny" ( [Darwin, 1859],Chapter 7). Despite Gregor Mendel being a contemporary of Darwin, the latter had probably no knowledge of the discoveries of the former [Galton, 2009, Sander, 1988]. Mendel’s idea of factors (genes) transmitting genetic information from one generation to the next [Mendel, 1866] came to popularity and great acceptance only several years after the death of Darwin in the year 1882. At Darwin’s time the modern knowledge of different gene expression or genomic imprinting as means to shape different morphologies were still unknown. Nevertheless, Darwin’s suggestion to resolve his major problem was that slight changes in a sterile worker could be passed to the next generations via its fertile parents ( [Darwin, 1859],Chapter 7). Thus, he was the first to think of a transfer of genetic information via closely related kin, even though he could not term it like this.
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