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Molecular Bases of Soft Coral Reproduction ResearchOnline@JCU This file is part of the following reference: Wessels, Wiebke (2016) Molecular bases of soft coral reproduction. PhD thesis, James Cook University. Access to this file is available from: https://researchonline.jcu.edu.au/51476/ The author has certified to JCU that they have made a reasonable effort to gain permission and acknowledge the owner of any third party copyright material included in this document. If you believe that this is not the case, please contact [email protected] and quote https://researchonline.jcu.edu.au/51476/ ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Australia Molecular Bases of Soft Coral Reproduction Dissertation submitted in partial fulfilment of the requirements for the degree of DOCTOR OF PHILOSOPHY in CORAL REEF STUDIES by Wiebke Wessels August 2016 DEDICATION To my family, who always believed in me I Statement of sources STATEMENT OF SOURCES I certify that the presented thesis Molecular Bases of Soft Coral Reproduction is, to the best of my knowledge and belief, original and my own work and has not been submitted in any form for another degree or diploma at any university or other institution of tertiary education. Information derived from the published or unpublished work of others has been acknowledged in the text and a list of references is given. Wiebke Wessels III Acknowledgements ACKNOWLEDGEMENTS First I want to thank my beloved family for the endless love and support that I received from my family throughout my studies. My advisory board, David Miller, for your guidance and patience with me and making this project turn out the best possible way. David Bourne, for answering all microbe−related questions that one can possibly have and improving this chapter so much. Susanne Sprungala, for your help, support and always open ears. Thank you for keeping me going and stay motivated. Sylvain Forêt, for answering all bioinformatics related questions and being patient when explaining the world of sequence analyses. Secondly I would like to thank former and present lab members and volunteers in the field… Eldon Ball, for setting me onto the (right) path of octocoral early development. Your excitement and interest in soft coral development is contagious and was most helpful during the late hours of coral spawning. Ira Cooke, for your great help, support and patience on the last bit of transcriptome and statistical analysis! Anthony Bertucci and Ben Mason, for being great boat drivers and diving buddies, for your endless help with my numerous field trips and staying calm when I was not anymore. Natalia Andrade, for coming along on almost every trip of mine, joining the ‘Team Softie’ and your humour ☺ Aurélie Moya, for your open door and time when I needed someone to get me back on track. Marta Espinheira, my field trips would have been a lot less successful without you and you made each field trip a memorable experience! Annie Bauer-Civiello and Georgina Torras-Jorda, for your amazing help during several coral spawning trips. V Acknowledgements Amin Esmail, Josh Mclntyre, Casey Chalkley, Kate Quigley for fruitful discussions in the lab or office and putting up with my endless carrot consume in the office ☺ For making my time in Australia unforgettable and adventurous… Laura Brannelly and Sarah Gierz for being there for me when I needed you most! And making life more fun! Elecia Johnston and David Care for making sure that I didn’t work 24ƒ7 and taking me out to see the sun every now and then and so much more. VI Statement of Contributions of Others STATEMENT OF CONTRIBUTIONS OF OTHERS Names, Titles and Affiliations Nature of Assistance Contribution of Co-Contributors Prof Dr David Miller, JCU Proposal writing Dr Susanne Sprungala, JCU Experimental design Dr Sue−Ann Watson, JCU Intellectual support Data collection Dr Susanne Sprungala, JCU Dr David Bourne, AIMS & JCU Data Analysis Dr Ira Cooke, JCU Microscopy Dr Eldon Ball, ANU Stipend JCU PRS Travel awards Robert Logan Fund Financial support GRS Graduate Scheme Research allocation CGC Student Award VII Thesis Abstract THESIS ABSTRACT The early development of the hard coral Acropora millepora has been described at both the morphological and molecular levels, and comparative studies are under way for other Scleractinia. Complementary data on Octocoral embryogenesis are, however, essential for a broader understanding of the different developmental strategies in Cnidaria and their evolution. Although the morphology of soft coral embryogenesis has been previously described, data are scarce and molecular data complementing the morphology are missing. This thesis involves the investigation of reproduction and early development in soft corals, using Lobophytum pauciflorum as model. As in the majority of soft corals, L. pauciflorum colonies are either female or male. The time of reproduction approximately coincides with the annual synchronised mass spawning of hard corals on the Great Barrier Reef (GBR), although in favourable years studied, it occurred a lunar month earlier. The process of gametogenesis is longer in the gonochoric soft coral than in the hermaphrodite hard coral A. millepora or the gonochoric hard coral Fungia concinna. However oogenesis is completed within a year, which is shorter than has been reported for a conspecific population in Taiwan. In addition, bidirectional sex change was observed in F. concinna, suggesting that this trait could also be a predominant reproductive feature in fungiid corals. After studying the gametogenic cycle of L. pauciflorum on the GBR and identifying the time point of spawning as 4 − 5 nights after the austral spring full moon, investigation of early development was possible. Following mixing of eggs and sperm, the first obvious signs of successful fertilisation occurred at the 16 − 32 cell stages. Through a series of cell divisions leading to irregular shapes, embryos developed into a swimming planula within 48 hours of fertilisation. Planulae then stayed in the water column for up to a week before settling and developing into a primary polyp with 8 tentacular buds in the 24 hours after settlement. Within the first week of metamorphosis, sclerites were visible inside the polyp tissue. Early development in L. pauciflorum appears to be slower than in A. millepora, and it was not possible to identify developmental stages corresponding to those of hard corals on the basis of morphology. Some similarities with hard corals were observed in terms of timing of expression of genes involved in calcification. In order to extend the knowledge on soft coral microbial communities, microbes associated with the entire colony were also investigated in Lobophytum pauciflorum. In a first approach the possibility of sex−specific differences in the core microbiome of L. pauciflorum IX Thesis Abstract was investigated. The profile of ribotypes present in at least 51 % of the samples was similar to the general profile when all ribotypes are considered. The microbial community in L. pauciflorum was dominated by Spirochaetes−related ribotypes, though a−Proteobacteria were the second most abundant class. Further, microbial profiles were very similar between male and female colonies in terms of highly abundant ribotypes, although sequences that cannot be assigned to any known bacteria were significantly more abundant among female than male colonies. Some sex−specific differences were detectable in ribotypes of low abundance. However, at class and family level, these differences do not persist, suggesting that the different ribotypes exhibit similar functions. The high abundance of unknown ribotypes provides the potential for discovery of novel bacterial strains with potentially different functions. In a second study, the effect of environmental stresses on the microbial community of L. pauciflorum was investigated. Microbial profiles of Lobophytum fragments exposed to 1) elevated temperatures, 2) reduced pH and 3) a combination of elevated temperatures and reduced pH for 12 days did not vary from control conditions. However, a clear pattern of colony specific microbial profiles at ribotype level emerged. A Spirochaete−dominated microbial community has only recently been identified in a gorgonian species in the Mediterranean Sea and deserves further characterisation. Combining the results of both studies showed that the microbial community of L. pauciflorum is conserved, with limited colony specific variability. Further, the microbial community appears robust to short term stresses, however the effect of longer−term stress deserves further investigation. X Table of Contents TABLE OF CONTENTS DEDICATION .............................................................................................................................. I STATEMENT OF SOURCES........................................................................................................ III ACKNOWLEDGEMENTS ........................................................................................................... V STATEMENT OF CONTRIBUTIONS OF OTHERS ....................................................................... VII THESIS ABSTRACT… ................................................................................................................. IX TABLE OF CONTENTS… ............................................................................................................ XI LIST OF FIGURES… ................................................................................................................
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