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THE UNIVERSITY OF HULL The behavioural function of pheromones in crayfish being a Thesis submitted for the Degree of Doctor of Philosophy in the University of Hull by Fiona Catherine Berry (MRes University of York; BSc University of St Andrews) September 2008 Contents Acknowledgements 1 Abstract 2 Chapter 1 - Introduction 3 t.l Biological communication 4 1.1.2 Chemical communication 4 1.1.3 Aquatic chemical signalling 6 1.1.4 Aquatic chemical signalling - crustaceans 7 1.2 The chemical nature of pheromones 9 1.2.1 Pheromone identification 9 1.2.2 Bioassay development 10 1.2.3 Chemical analysis 11 1.3 The use of pheromones to control invasive species 11 1.4 Biological and ecology of crayfish (The Astacoidea) 12 1.4.1 Morphology and growth 13 1.4.2 Life history strategies 14 1.4.3 Crayfish in the UK and Europe 15 1.4.3.1 White-clawed crayfish in the UK 16 1.4.3.2 Non-indigenous crayfish 17 1.4.4 Controlling invasive crayfish species 19 1.5 Current knowledge of the role of chemical communication in crayfish, in the context of behaviour 21 1.5.1 Inter-specific chemical communication 22 1.5.2 Reproduction 22 1.5.3 Mate choice 25 1.5.4 Agonism 26 1.6 Aims and Objectives 27 Chapter 2 - General methodology 31 2.1 Animal maintenance 32 2.1.1 Collection 32 2.1.2 Legislation and holding 32 2.2 Experimental procedures 33 2.2.1 Identification 33 2.2.2 Isolation 33 2.2.3 Blindfolding 34 2.2.4 General experimental protocols 34 2.2.5 Checking for receptivity 34 2.2.6 Urine visualisation 35 2.2.7 Urine sampling technique 36 2.2.8 Conditioned water 36 2.2.9 Urine blocking 36 2.2.10 Behavioural analysis 37 2.2.11 Physiological measurements 37 2.2.11.1 Heartbeat analysis 3 7 2.2.11.2 Scaphognathite beat analysis 39 2.2.12 Statistical analysis 43 Chapter 3 - Urinary signalling in crayfish social interactions: miscommunication between females and males 44 3.1 Introduction 45 3.2 Materials and methods 47 3.2.1 Animals 47 3.2.2 Urinary signalling in social interactions 48 3.2.3 Urine visualisation procedure 48 3.2.4 Social interactions 48 3.2.5 Behavioural analysis 49 3.2.6 Agonistic behaviours 50 3.2.7 Reproductive behaviours 51 3.2.8 Urine release analysis 51 3.2.9 Statistical analysis 51 3.3 Results 52 3.3.1 Sex-specific differences in chemical signalling 52 3.3.2 Mating behaviour 53 3.3.3 Female urine release in sexual interactions 54 3.3.4 Agonistic behaviour 55 3.3.5 Urine release in agonistic interactions 55 3.4 Discussion 56 3.4.1 The role of female urinary cues in eliciting male courtship 56 iii 3.4.2 Absence of male urine release during courtship 57 3.4.3 Sexual conflict and urine signalling 57 3.4.4 Urinary signalling in agonistic interactions 58 3.4.5 The'dilemma'of urine signalling 58 3.4.5 Conclusions 59 Chapter 4 - Mate choice in crayfish, Pacifastacus leniusculus: Do females choose dominant partners when given a free choice? 60 4.1 Introduction 61 4.2 Materials and methods 63 4.2.1 Animal maintenance 63 4.2.2 Fight interactions 64 4.2.3 Mate choice experiments 64 4.2.4 Mate choice analysis 65 4.2.5 Statistical analysis 66 4.3 Results 66 4.4 Discussion 67 Chapter 5 - Ventilatory and heartbeat activity of crayfish, Pacifastacus leniusculus, during reproductive behaviours 70 5.1 Introduction 71 5.2 Materials and methods 72 5.2.1 Animal holding conditions 72 5.2.2 Mating interactions 73 5.2.3 Physiological recordings 73 5.2.4 Physiological analysis 74 5.2.5 Statistical analysis 74 5.3 Results 74 5.4 Discussion 77 Chapter 6 - Ventilatory measures in crayfish during social interactions: the effects of social dominance and urinary signalling 80 6.1 Introduction 81 6.2 Materials and methods 82 iv 6.2.1 Animal maintenance 82 6.2.2 Agonistic interactions 83 6.2.3 Ventilation rate recording 84 6.2.4 Urine visualisation procedure in P. clarkii 84 6.2.5 Behavioural analysis 85 6.2.6 Ventilation rate analysis 85 6.2.7 Statistical analysis 85 6.3 Results 86 6.4 Discussion 91 Chapter 7 - Development of behavioural and physiological assays to assess discrimination of male and female odours in crayfish, Pacifastacus lenimculus 94 7.1 Introduction 95 7.2 Materials and methods 97 7.2.1 Animal holding conditions 97 7.2.2 Urine and conditioned water collection 98 7.2.3 General bioassay procedure 98 7.2.4 Behavioural bioassay 99 7.2.5 Heartbeat assay 100 7.2.6 Ventilation assay 102 7.3 Results 102 7.3.1 Behavioural bioassay 102 7.3.2 Heartbeat assay 105 7.3.3 Ventilation assay 107 7.4 Discussion 108 7.4.1 Behavioural bioassay 109 7.4.2 Physiological assays 110 Chapter 8 - The effects of artificial urine release on the fight dynamics of crayfish, Pacifastacus leniusculm 113 8.1 Introduction 114 8.2 Materials and methods 117 8.2.1 Animal holding conditions 117 8.2.2 Urine blocking 117 8.2.3 General fight procedure 118 8.2.4 Stimulus collection and preparation 119 8.2.5 Stimulus introduction in fight two 119 8.2.6 Behavioural analysis 120 8.2.7 Statistical analysis 121 8.3 Results 121 8.4 Discussion 123 8.4.1 The effect of dominant urinary signals in fights 123 8.4.2 Candidate substances for crayfish status cues 124 Chapter 9 - Discussion 126 9.1 Chemical communication in crayfish 127 9.1.1 Chemical signalling during mating 127 9.1.2 Aggressive chemical signalling 129 9.1.3 Signal delivery 130 9.2 Future research 130 9.2.1 Determination of the chemical nature of crayfish pheromones 130 9.2.1.1 Sex pheromones 13 0 9.2.1.2 Dominance pheromones 132 9.2.2 Application of methodologies 132 9.2.3 Mate choice studies 133 9.3 Conclusion 134 References 135 VI Figures 1.1 Ventral views of the crayfish Procambarus clarkii (A) Form I male (B) Female 14 1.2 Distribution of white-clawed crayfish in the UK (A) 1979-1988(6) 1999-2008 16 1.3 Distribution of signal crayfish in the UK (A) 1969 - 1978 (B) 1979-1988 (C) 1989 - 1998 (D) 1999 - 2008 19 2.1 Blindfolded signal crayfish 34 2.2 Experimental equipment used to measure physiological parameters (heart rate, ventilation rate) of crayfish 38 2.3 Infra-red sensors used to measure (A) heartbeat activity (B) scaphognathite movement of crayfish 39 2.4 Ventilation rate recordings of male signal crayfish (A) right scaphognathite recording (B) left scaphognathite recording 41 2.5 Ventilation rate recording of an isolated male red swamp crayfish engaged in fighting (A) right scaphognathite recording (B) left scaphognathite recording 42 3.1 Urine release by dominant and subordinate crayfish in social interactions 53 3.2 Ethogram of a sexual interaction between signal crayfish 54 3.3 Urine release by female crayfish during sexual interactions 55 4.1 Experimental set up from mate choice experiments (A) acclimation period (B) experimental period 65 5.1 Ventilation rate of male and female crayfish during reproductive behaviours 75 5.2 Scaphognathite activity of (A) female and (B) male signal crayfish during reproductive behaviours 76 5.3 Heart rate of female crayfish during reproductive interactions 77 6.1 Scaphognathite activity of male red swamp crayfish during aggressive fighting 87 6.2 Ventilation rate of crayfish during agonistic behaviours (A) Procambarus clarkii (B) Pacifastacus leniusculus 88 6.3 Scaphognathite activity of male signal crayfish at the onset of aggressive fighting 89 vii 6.4 Ventilation rate of male red swamp crayfish during urinary signalling and aggressive fighting 90 6.5 Scaphognathite activity of male red swamp crayfish at the onset of urinary signalling 90 7.1 Number of male crayfish which mounted a female dummy following introduction of conspecific odours 103 7.2 Timed behavioural responses of male crayfish to a female dummy (A) mounting the female dummy (B) touching the female dummy with the chelae 104 7.3 Heart rate recording for male crayfish following the release of female urine 105 7.4 The number of male crayfish showing cardiac arrest to introduced stimuli 106 7.5 Standardised heart rate recorded by male crayfish following stimulus release 107 7.6 Number of male crayfish displaying ventilatory arrest to introduced stimulus 108 8.1 Fight dynamics of crayfish with unblocked nephropores on day one and blocked nephropores on day two (A) mean fight duration (B) mean fight intensity 122 Tables 1.1 The life history characteristics of three crayfish found in Europe 15 1.2 Native and non-indigenous crayfish of Europe 17 3.1 Agonistic and sexual behaviours displayed by signal crayfish in social interactions 50 3.2 Two-way repeated measures ANOVA results for urine release during social interactions 52 4.1 Mate choice and behaviour among female crayfish 67 5.1 Two-way repeated measures ANOVA results for ventilation rate of male and female crayfish during reproductive behaviours 75 6.1 Two-way repeated measures ANOVA results for the ventilation rate of male crayfish during agonistic behaviours (A) signal crayfish 86 (B) red swamp crayfish 87 8.1 Two-way repeated measures ANOVA results for (A) fight intensity (B) fight duration of second day fights where crayfish had blocked nephropores and were exposed to different experimental conditions 121 viii Acknowledgements I would firstly like to thank my supervisor, Thomas Breithaupt, for his enthusiasm towards my research and for the many hours spent discussing project ideas.
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  • Arrival of Non-Indigenous Crayfish Species

    Arrival of Non-Indigenous Crayfish Species

    Knowl. Manag. Aquat. Ecosyst. 2016, 417, 37 Knowledge & © G. Kotovska et al., Published by EDP Sciences 2016 Management of Aquatic DOI: 10.1051/kmae/2016024 Ecosystems www.kmae-journal.org Journal fully supported by Onema RESEARCH PAPER East European crayfish stocks at risk: arrival of non-indigenous crayfish species Ganna Kotovska1, Dmytro Khrystenko1,2,*,Jiří Patoka3,a and Antonín Kouba4,a 1 Institute of Fisheries of the National Academy of Agrarian Sciences of Ukraine, Obuhivska 135, Kyiv 03164, Ukraine 2 Cornell University, College of Agriculture and Life Sciences, International Programs, B75 Mann Library, Ithaca, NY 14853, USA 3 Czech University of Life Sciences Prague, Faculty of Agrobiology, Food and Natural Resources, Department of Zoology and Fisheries, Kamycká 129, Prague 6–Suchdol 165 21, Czech Republic 4 University of South Bohemia in České Budějovice, Faculty of Fisheries and Protection of Waters, South Bohemian Research Center of Aquaculture and Biodiversity of Hydrocenoses, Zátisí 728/II, Vodňany 389 25, Czech Republic Abstract – An increasing number of non-indigenous crayfish species (NICS) of apparently pet trade origin have become established particularly in Europe. Especially alarming are recent confirmation of two distantly separated marbled crayfish Procambarus fallax f. virginalis populations in Ukraine and indications of more North American cambarids present in the local pet market. The present study aimed to investigate crayfish species availability within the Ukrainian pet trade together with the climate match and risk they represent to the freshwater ecosystems generally and indigenous crayfish species in particular. Altogether, 15 NICS belonging to all three crayfish families were detected. Considering their origin, availability, probability of establishment, invasiveness and further aspects, marbled crayfish and red swamp crayfish Procambarus clarkii appear to be potentially the most troubling.
  • Durham E-Theses

    Durham E-Theses

    Durham E-Theses The Inuence of Red Colouration on Human Perception of Aggression and Dominance in Neutral Settings WIEDEMANN, DIANA How to cite: WIEDEMANN, DIANA (2016) The Inuence of Red Colouration on Human Perception of Aggression and Dominance in Neutral Settings, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/11866/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk 2 The Influence of Red Colouration on Human Perception of Aggression and Dominance in Neutral Settings ___________________________________________________________________________ Diana Wiedemann Thesis submitted for the degree of Doctor of Philosophy Department of Anthropology, Durham University January 2016 i Abstract Abstract For both humans and nonhuman species, there is evidence that red colouration signals both emotional states (arousal/anger) and biological traits (dominance, health, and testosterone). The presence and intensity of red colouration correlates with male dominance and testosterone in a variety of animal species, and even artificial red stimuli can influence dominance interactions.
  • Better Wear Red? the Influence of the Color of Sportswear on the Outcome

    Better Wear Red? the Influence of the Color of Sportswear on the Outcome

    Submitted by Matthias Nikolaus Hilgarth, BSc Submitted at Department of Economics Supervisor Dr. Mario Lackner October 2020 Better wear red? The influence of the color of sportswear on the outcome of Olympic sport competitions Master Thesis to obtain the academic degree of Master of Science in the Master’s Program Economics JOHANNES KEPLER UNIVERSITY LINZ Altenbergerstraße 69 4040 Linz, Österreich www.jku.at DVR 0093696 Sworn Declaration I, Matthias Nikolaus Hilgarth, hereby declare under oath that the thesis submitted is my own unaided work, that I have not used sources other than the ones indicated, and that all direct and indirect sources are acknowledged as references. This printed thesis is identical with the electronic version submitted. Linz, Place and Date Matthias Nikolaus Hilgarth 2 "All I am or can be I owe to my angel mother." Abraham Lincoln For Mom 3 Acknowledgments First and foremost, I would like to express my deepest gratitude to Dr. Mario Lackner for providing me with the topic of this Master’s thesis, for the trust in me to work independently, the patience to give me time and support whenever I needed it. Moreover, I am very grateful to Alexander Ahammer, PhD for his assistance in the whole process. Additionally I would like to thank Univ.-Prof. Dr. Martin Halla for his valuable suggestions. Thanks the whole Department of Economics for providing such a an enjoyable environment for learning and working. Your efforts to make students part of the department are truly appreciated. A special thanks to my partner Martina, for your kindness and encouragement.
  • Development of New Genome-Informed Genotyping Tools for Aphanomyces Astaci

    Development of New Genome-Informed Genotyping Tools for Aphanomyces Astaci

    Development of new genome-informed genotyping tools for Aphanomyces astaci Submitted by Diana Minardi to the University of Exeter as a thesis for the degree of Doctor of Philosophy in Biological Sciences In May 2017 This thesis is available for Library use on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I certify that all material in this thesis which is not my own work has been identified and that no material has previously been submitted and approved for the award of a degree by this or any other University. Signature: ………………………………………………………….. 1 Acknowledgements I would like to sincerely thank my supervisors Dr Mark van der Giezen, Dr David Studholme, and Dr Birgit Oidtmann for their invaluable guidance and support during these 3 years (plus a bit) of PhD, both practically and in the writing of the thesis. I would like to thank Cefas and the University of Exeter for funding the project and for giving me the opportunity to embrace this challenge and adventure. I would like to thank all of the University of Exeter “Biocat” lab and office, with whom I’ve spent most of my coffee time. But especially Mirella (for the laughs and cries) and Simone (for the sweaty runs around campus). I would also like to thank Chiara for her constant support, encouragement, and beer. Finally, I would like to thank Tony, for his patience, and my family, for their “overseas” support: I’ve finally finished “the shrimp” book! 2 Abstract Aphanomyces spp. are water moulds, eukaryotic fungus-like organisms, belonging to the class Oomycota.