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Variation in Male Post-Copulatory Investment: Ontogeny to Progeny

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Variation in Male Post-Copulatory Investment: Ontogeny to Progeny ERIN L MACARTNEY VARIATION IN MALE POST-COPULATORY INVESTMENT: ONTOGENY TO PROGENY A thesis in fulfilment of the requirements for the degree of Doctor of Philosophy September 2019 School of Biological, Earth and Environmental Sciences Faculty of Science THESIS SHEET Surname/Family Name : Macartney Given Name/s : Erin Larissa Abbreviation for degree as give in the University calendar : PhD Faculty : Science School : School of Biological, Earth, and Environmental Sciences Thesis Title : Variation in male post-copulatory investment: ontogeny to progeny Condition-dependence theory has been widely applied to exaggerated male signaling traits, and more recently to certain sperm and semen traits. However, such post-copulatory traits can be highly cryptic and multi-faceted, and the published literature shows considerable variation in trait expression due to male condition. This thesis aims to uncover sources of variation in post-copulatory trait expression, and to incorporate male investment in ejaculate and offspring quality into a condition-dependence life-history framework. Using meta-analyses and meta-regression in Chapter Two, I demonstrate that much of the variation in post-copulatory responses to nutrient limitation (used as a manipulation of condition) is accounted for by differences in the type of nutrients, the ontogenetic life-stage when nutrients are limited, and the type of trait. Trait responses are also taxon-specific. In Chapter Three, I empirically demonstrate that developmental nutrient limitation strongly reduces testes and accessory gland size, as well as sperm movement within the female reproductive tract, and that adult diet does not alter such responses in the neriid fly, Telostylinus angusticollis. In Chapters Four and Five, I consider condition-dependent effects on offspring quality. In Chapter Four, I argue that non-genetic paternal effects conferred through epigenetic factors may also be costly, condition-dependent traits. In Chapter Five, I use T. angusticollis to test if frequent mating results in a condition-dependent reduction in fecundity and offspring quality. Surprisingly, frequent mating did not result in reduced fecundity or offspring quality, but did result in a reduced mating rate. Finally, in Chapter Six, I demonstrate that male Drosophila melanogaster suffer sperm depletion across successive matings, but the rate of depletion is not dependent on diet or genotype. Instead, males vary in ability to mate multiply, and individuals that mate more also transfer more sperm. Thus, variation in male post-copulatory performance depends substantially on unknown factors that are unrelated to nutrition and genotype. Overall, this thesis demonstrates that sperm and semen traits are affected by many different factors, and that variation in such traits can be highly complicated. Understanding differences in post- copulatory trait expression will provide increased understanding into the evolution of mating systems and reproductive investment. Declaration relating to disposition of project thesis/dissertation I hereby grant to the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or in part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstracts International (this is applicable to doctoral theses only). 20/09/2019 …………………………………………………………… ……………………………………..……………… ……….……………………...…….… Signature Witness Signature Date The University recognises that there may be exceptional circumstances requiring restrictions on copying or conditions on use. Requests for restriction for a period of up to 2 years must be made in writing. Requests for a longer period of restriction may be considered in exceptional circumstances and require the approval of the Dean of Graduate Research. Date of completion of requirements for FOR OFFICE USE ONLY Award: ORIGINALITY STATEMENT ‘I hereby declare that this submission is my own work and to the best of my knowledge it contains no materials previously published or written by another person, or substantial proportions of material which have been accepted for the award of any other degree or diploma at UNSW or any other educational institution, except where due acknowledgement is made in the thesis. Any contribution made to the research by others, with whom I have worked at UNSW or elsewhere, is explicitly acknowledged in the thesis. I also declare that the intellectual content of this thesis is the product of my own work, except to the extent that assistance from others in the project's design and conception or in style, presentation and linguistic expression is acknowledged.’ S.. …………………………………………….............. 20/09/2019 …………………………………………….............. INCLUSION OF PUBLICATIONS STATEMENT UNSW is supportive of candidates publishing their research results during their candidature as detailed in the UNSW Thesis Examination Procedure. Publications can be used in their thesis in lieu of a Chapter if: • The student contributed greater than 50% of the content in the publication and is the “primary author”, i.e., the student was responsible primarily for the planning, execution and preparation of the work for publication • The student has approval to include the publication in their thesis in lieu of a Chapter from their supervisor and Postgraduate Coordinator. • The publication is not subject to any obligations or contractual agreements with a third party that would constrain its inclusion in the thesis Please indicate whether this thesis contains published material or not. This thesis contains no publications, either published or submitted for ☐ publication Some of the work described in this thesis has been published and it has been ☒ documented in the relevant Chapters with acknowledgement This thesis has publications (either published or submitted for publication) ☐ incorporated into it in lieu of a chapter and the details are presented below CANDIDATE’S DECLARATION I declare that: • I have complied with the Thesis Examination Procedure • where I have used a publication in lieu of a Chapter, the listed publication(s) below meet(s) the requirements to be included in the thesis. Name Signature Date (dd/mm/yy) Erin L Macartney 10/01/2020 COPYRIGHT STATEMENT ‘I hereby grant the University of New South Wales or its agents a non-exclusive licence to archive and to make available (including to members of the public) my thesis or dissertation in whole or part in the University libraries in all forms of media, now or here after known. I acknowledge that I retain all intellectual property rights which subsist in my thesis or dissertation, such as copyright and patent rights, subject to applicable law. I also retain the right to use all or part of my thesis or dissertation in future works (such as articles or books).’ ‘For any substantial portions of copyright material used in this thesis, written permission for use has been obtained, or the copyright material is removed from the final public version of the thesis.’ Signed ……………………………………………........................... Date ……………………………………………10/01/2020 . ............................. AUTHENTICITY STATEMENT ‘I certify that the Library deposit digital copy is a direct equivalent of the final officially approved version of my thesis.’ Signed ……………………………………………........................... Date ………………………10/01/202………………… 0 …................... PUBLICATIONS, PRESENTATIONS AND AWARDS Publications Macartney, E.L., Crean A. J., Nakagawa, S., Bonduriansky, R. (2019). Effects of nutrient limitation on sperm and seminal fluid: A systematic review and meta-analysis. Biological Reviews: 94, 1722—1739. Macartney E.L., Nicovich R. R., Bonduriansky R., Crean A. J. (2018). Developmental diet irreversibly shapes male post-copulatory traits in the neriid fly (Telostylinus angusticollis). Journal of Evolutionary Biology: 31, 1894–1902. Macartney E.L., Crean A.J., Bonduriansky R. (2018). Epigenetic paternal effects as costly, condition-dependent traits. Hereditary: 121, 248–256. Macartney E.L., Crean A.J., Bonduriansky R. (2017). Adult dietary protein has age- and context-dependent effects on male post-copulatory performance. Journal of Evolutionary Biology: 38, 42–49. Nicovich, P.R., Macartney, E.L., Whan, R.M. and Crean, A.J. (2015). Measuring sperm movement within the female reproductive tract using Fourier analysis. Microscopy and Microanalysis: 21, 256–263. Presentations UNSW 3-Minute Thesis Competition, Sydney, Australia. (2018). You are what you eat: dietary effects on male reproduction. Science Faculty Research Show Case, Sydney, Australia. (2018). You are what you eat: dietary effects on male reproduction – first place. Australasian Evolutionary Society conference, Hobart, Australia. (2017). Condition-dependent post-copulatory traits: a meta-analysis. Biology of Spermatozoa Conference, Bakewell, United Kingdom. (2017). Cryptic non-genetic paternal effects as condition-dependent traits. European Evolutionary Society conference, Groningen, Netherlands. (2017). Cryptic non-genetic paternal effects as condition-dependent traits. Behaviour conference, Cairns, Australia. (2015). No cost of mating for male neriid flies? International Society for Behavioural Ecology
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