EXPERIMENTAL APPROACHES TO UNRAVEL PROXIMATE MECHANISMS OF PARASITOID SEARCHING AND PATCH LEAVING BEHAVIOUR DOCTORAL THESIS by JENNIFER M. UHLIG-HERRMANN BREMEN, OCTOBER 2015 EXPERMENTELLE ANSÄTZE ZUM AUFDECKEN PROXIMATER MECHANISMEN DES SUCH- UND AUFGABEVERHALTENS VON PARASITOIDEN DISSERTATION ZUR ERLANGUNG DES AKADEMISCHEN GRADES DOKTOR DER NATURWISSENSCHAFTEN ‐ Dr.rer.nat.‐ VORGELEGT IM FACHBEREICH 2 ‐ BIOLOGIE/CHEMIE, UNIVERSITÄT BREMEN von JENNIFER M. UHLIG-HERRMANN BREMEN, OKTOBER 2015 Examination committee/ Prüfungsausschuss 1st Reviewer/ Erstgutachter: Prof. Dr. Thomas S. Hoffmeister Population and Evolutionary Ecology Group, FB2 Biology/Chemistry, University of Bremen, Germany 2nd Reviewer/ Zweitgutachter: Prof. Dr. Dorothea Eisenhardt Neurobiology, Department of Biology, Chemistry, Pharmacy, Freie Universität Berlin, Germany 1st Additional Examiner/ 1. Prüfer: Prof. Dr. Christian Wild Marine Ecology, Biology/Chemistry, University of Bremen, Germany 2nd Additional Examiner/ 2. Prüfer: Prof. Dr. Martin Diekman Vegetation Ecology and Conservation Biology, Institute of Ecology, FB 2, University of Bremen, German 3 Contents Table of Contents Examination committee/ Prüfungsausschuss ......................................................... 3 Summary .............................................................................................................. 7 Zusammenfassung ................................................................................................ 9 1 General introduction and overview .................................................................... 11 I.2 Study organisms .................................................................................................. 17 1.3 References .......................................................................................................... 20 Chapter 2- Analysis of walking behavior as a correlate of motivation in two parasitic wasps .................................................................................................... 25 2.1 Abstract: ............................................................................................................. 26 2.2 Introduction........................................................................................................ 26 2.3 Material and Methods........................................................................................ 29 2.3.1 Venturia canescens ...................................................................................... 30 2.3.2 Leptopilina heterotoma ............................................................................... 30 2.3.3 Statistical Analysis ....................................................................................... 32 2.4 Results ................................................................................................................ 33 2.4.1 Patch leaving and movement in Venturia canescens ................................. 33 2.4.2 Patch leaving and movement in Leptopilina heterotoma ........................... 40 2.5 Discussion ........................................................................................................... 46 2.5.1 Response of V. canescens ............................................................................ 47 2.5.2 Response of L. heterotoma ......................................................................... 48 2.6 References .......................................................................................................... 51 Chapter 3 - Effects of octopamine-feeding on responsiveness to the patch in the parasitoid Leptopilina heterotoma ....................................................................... 55 3.1 Abstract .............................................................................................................. 56 3.2 Introduction........................................................................................................ 56 3.3 Materials and methods ...................................................................................... 59 3.3.1 Study species ............................................................................................... 59 3.3.2 General protocol ......................................................................................... 59 3.3.3. Drug treatment .......................................................................................... 60 3.3.4 Experimental approach ............................................................................... 60 3.3.5 Behavioural scoring ..................................................................................... 61 3.3.6 Data Analysis ............................................................................................... 61 3.4 Results ................................................................................................................ 62 4 Contents 3.4.1 Effects of OA treatment on latency ............................................................. 62 3.4.2 Effects of OA treatment on PRT and on-patch behaviour .......................... 63 3.5 Discussion ........................................................................................................... 64 3.6 References .......................................................................................................... 69 Chapter 4 - Octopamine affects the dynamic response to danger in a parasitoid wasp ................................................................................................................... 74 4.1 Abstract .............................................................................................................. 75 4.2 Introduction........................................................................................................ 75 4.2.1 Dealing with risk .......................................................................................... 75 4.2.2 Age ............................................................................................................... 76 4.2.3 Neuronal modulation of decision making ................................................... 76 4.3 Materials and Methods ...................................................................................... 78 4.3.1 General experimental set up ....................................................................... 78 4.3.2 Pre-experimental treatment ....................................................................... 79 4.3.3 Feeding treatment....................................................................................... 79 4.3.4 Data analysis ................................................................................................ 80 4.4 Results ................................................................................................................ 80 4.5 Discussion ........................................................................................................... 82 4.6 References .......................................................................................................... 86 Chapter 5 - The effects of biogenic amines on intrasexual competition and longevity in female Asobara tabida ..................................................................................... 91 5.1 Abstract .............................................................................................................. 92 5.2 Introduction........................................................................................................ 92 5.3 Material and Methods........................................................................................ 95 5.3.1 Behavioural observations ............................................................................ 95 5.3.2 Effects of intrasexual competition and aminergic treatment on longevity 97 5.4.1 Behavioural observations ............................................................................ 98 5.4.2 Effects of aminergic treatment on longevity ............................................ 101 5.5 Discussion ......................................................................................................... 102 5.6 References ........................................................................................................ 108 Chapter 6 - The effects of biogenic amines on intrasexual competition and longevity in males of the parasitic wasp Asobara tabida .................................................... 114 6.1 Abstract ............................................................................................................ 115 6.2 Introduction...................................................................................................... 115 6.3 Methods ........................................................................................................... 117 6.3.1 Intrasexual conflicts and behaviour .......................................................... 117 5 Contents 6.3.2 Effects of amine -treatment and competition on longevity .............. 119 6.4 Results ........................................................................................................ 120 6.4.1 Intrasexual conflicts and behaviour .......................................................... 120 6.4.3 Effects of amine treatment and competition on longevity ................ 123 6.5 Discussion ......................................................................................................... 124 7 General discussion and conclusion ..................................................................