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Bacteria on Coralline Algae and Their Role As Sea Urchin Settlement Cues

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Bacteria on Coralline Algae and Their Role As Sea Urchin Settlement Cues Bacteria on coralline algae and their role as sea urchin settlement cues Shaun Nielsen A thesis in fulfilment of the requirements for the degree of Doctor of Philosophy School of Biological, Earth and Environmental Sciences Faculty of Science 2013 THE UNIVERSITY OF NEW SOUTH WALES - Thesis/Dissertation Sheet Surname or Family name:Nielsen First name:Shaun Other name/s: Abbreviation for degree as given in the University calendar: PhD School:Biological, Earth and Environmental Sciences Faculty:Science Title:Bacteria on coralline algae and their role as sea urchin settlement cues Abstract Most benthic marine invertebrates have a biphasic life cycle, in which a planktonic larval stage alternates with a benthic ad ult stage. The transition between the larval and adult stage is typically guided by habitat-derived settlement cues and thus understanding the nature and distribution of settlement cues is a central theme in larval ecology. Both coralline algae and their epiphytic bacterial biofilms can be important settlement cues for marine invertebrate larvae, but the relationship between settlement and specific communities of bacteria is largely unknown. I investigated bacterial mediated settlement for larvae of the Australian sea urchins Heliocidaris erythrogramma and Holopneustes purpurascens and compared this to the community ecology of bacteria on coralline algae. I conducted a meta-analysis of putative larval cues from macroalgae to test the importance of coralline algae as settlement cues for invertebrate larvae generally and sea urchin larvae specifically. The meta-analysis revealed that coralline algae were the most inductive macroalgae for a variety of larval groups, but epiphytic bacteria only enhanced larval settlement for a few larval groups including sea urchins. Using larvae of H. erythrogramma and H. purpurascens, I next showed in larval settlement assays that bacteria on coralline algae enhanced settlement for both species but only larvae of H. erythrogramma responded to specific variation in the bacterial community composition. This specificity of response was then tested by isolating bacteria from the surface of coralline algae, and testing these against settlement by both sea urchins. One bacterium, Pseudoalteromonas luteoviolacea, was isolated from different species of corallines and induced larval settlement of both sea urchins, suggesting a common settlement cue across coralline algae. 16S rRNA tag sequencing surveys of the relative abundance of bacteria on these algae in the field indicated a high abundance of bacterial groups not examined in larval assays and an extremely low number of the genus Pseudoalteromonas, suggesting a very low abundance of P. luteoviolacea in natural communities. In summary, this thesis presents a systematic overview of larval settlement studies using meta -analysis, a statistical framework for correlating larval settlement and bacterial communities and provides an understanding of dominant bacterial members that have yet to be examined in larval settlement studies. Declaration relating to disposition of project thesis/dissertation I hereby grant to the Univ ersity of New South Wales or its agents the right to archiv e and to make av ailable my thesis or dis sertation in whole or in part in the Univ ersity libraries in all f orms of media, now or here af ter known, subject to the prov isions of the Copy right Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in f uture works (such as articles or books) all or part of this thesis or dissertation. I also authorise Univ ersity Microf ilms to use the 350 word abstract of my thesis in Dissertation Abstracts International (this is applicable to doctoral theses only ). …………………………………………………………… ……………………………………..……………… ………...…….… Signature Witness 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. FOR OFFICE USE ONLY Date of completion of requirements for 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.’ Signed ................................................................. Date .................................................................... COPYRIGHT STATEMENT ‘I hereby grant the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or 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 proprietary 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 Abstract International (this is applicable to doctoral theses only). I have either used no substantial portions of copyright material in my thesis or I have obtained permission to use copyright material; where permission has not been granted I have applied/will apply for a partial restriction of the digital copy of my thesis or dissertation.' Signed ……………………………………………........................... Date ……………………………………………........................... AUTHENTICITY STATEMENT ‘I certify that the Library deposit digital copy is a direct equivalent of the final officially approved version of my thesis. No emendation of content has occurred and if there are any minor variations in formatting, they are the result of the conversion to digital format.’ Signed ……………………………………………........................... Date ……………………………………………........................... Table of contents Abstract ........................................................................................................................................I Chapter 1 ..................................................................................................................................... 1 1.1 The larval stage of benthic marine invertebrates ....................................................... 1 1.2 Larval settlement cues ............................................................................................. 2 1.3 Coralline algae as settlement cues ............................................................................ 4 1.4 Bacteria on coralline algae as settlement cues ........................................................... 5 1.5 Understanding bacterial settlement cues .................................................................. 6 1.6 Study organisms ...................................................................................................... 7 1.6.1 Sea urchins .......................................................................................................... 7 1.6.2 Coralline algae ..................................................................................................... 9 1.7 Thesis aims .............................................................................................................. 9 1.8 Chapter synopsis.................................................................................................... 10 Chapter 2 ................................................................................................................................... 12 2.1 Introduction ............................................................................................................. 12 2.2 Methods .................................................................................................................. 15 2.2.1 Database searching ............................................................................................ 15 2.2.2 Experimental criteria .......................................................................................... 15 2.2.3 Meta-analysis predictor variables ........................................................................ 16 2.2.3 Statistical analysis .............................................................................................. 17 2.3 Results ..................................................................................................................... 19 2.3.1 Overall larval settlement across all substrata ....................................................... 19 2.3.2 Effect of latitude ................................................................................................ 19 2.3.3 Effect of larval time to competence ..................................................................... 19 2.3.4 Effect of experimental conditions .......................................................................
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