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The Role of Turbulence in Broadcast Spawning and Larval THE ROLE OF TURBULENCE IN BROADCAST SPAWNING AND LARVAL SETTLEMENT IN FRESHWATER DREISSENID MUSSELS A Thesis Presented to The Faculty of Graduate Studies of The University of Guelph by NOEL PETER QUINN In partial fulfilment of requirements for the degree of Doctor of Philosophy December, 2009 © Noel P. Quinn, 2009 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de ('edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 OttawaONK1A0N4 Canada Canada Your file Voire reference ISBN: 978-0-494-58276-3 Our file Notre reference ISBN: 978-0-494-58276-3 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. 1*1 Canada ABSTRACT THE ROLE OF TURBULENCE IN BROADCAST SPAWNING AND LARVAL SETTLEMENT IN FRESHWATER DREISSENID MUSSELS Noel Peter Quinn Advisor: University of Guelph, 2009 Professor J. D. Ackerman The role of turbulence has been shown theoretically to influence external fertilization and larval settlement/transport in benthic invertebrates. This is especially true for turbulence generated by bottom roughness in the near-bed region of lakes. This thesis examined the role of bottom roughness created by the presence of freshwater mussels (Dreissena polymorpha and D. bugensis) by addressing three objectives: (1) how fertilization success is influenced by water velocity and near-bed turbulence generated by bottom roughness in a laboratory flow chamber and in the field (Evans Point, Lake Erie); (2) how larval transport and settlement is influenced by this near-bed turbulence in laboratory flow chamber and field (Lake Erie) experiments; and (3) how gamete and larval transport are influenced by the ratio of roughness spacing (X) to roughness height (k) using computational fluid dynamic (CFD) modeling. Results indicated that dreissenid mussels are sperm limited, but the extent to which sperm dilution affects them is lower than what has been reported for other broadcast spawners. The nature of near-bed turbulence was determined through digital particle imaging velocimetry (PIV) measurements in the laboratory and acoustic Doppler velocimetry (ADV) measurements in the field. Laboratory results indicated the importance of bottom roughness on the flow regime, notably skimming flow was observed in the high density mussel configuration and wake interference was observed in the mussel patch configuration. Fertilization success and larval settlement in the field was highest under the mussel patch configuration, which was positively associated with turbulent ejections. Results from the CFD modeling, which incorporated a released scalar as a proxy for gamete and larvae, accurately predicted the flow regimes classified using the ratio of X/k (e.g., < 8 for skimming flow and ~8 for wake interference flow), but this varied with the geometry of the modeled roughness elements. These results indicate that the spatial configuration of bottom roughness, including mussels, determines the flow regime (i.e., skimming vs. wake interference flow), which in turn affects fertilization success and larval transport/settlement in benthic species. ACKNOWLEDGEMENTS I would like to thank Kelly McNichols, Brendan Hunt, Sarah Glover, Peter Blouw, Rob Schindler, and Neil Menezes for all the assistance in the laboratory and field. I would also like to thank Greg Nishihara and Patrick Ragaz for their assistance with the flume and,PIV setup. Throughout my PhD, I have had invaluable feedback from my PhD committee, so I would like to thank Dr. David Barton, Dr. Ray Kostaschuk, and Dr. KevinJVlcCann for helping me over these last few years. I would also like to thank Dr. Pete Jumars for acting as my External Reviewer, and providing some invaluable insight into my thesis. My advisor, Dr. Joe Ackerman, has made me a better researcher, writer, and scientist, and without his guidance and friendship I would not be where I am today. I would to thank my parents for all their support over the last few years, and finally I would like to thank my wife, Tara, for being there for me when I needed someone to bounce ideas off of, when I needed guidance, or even when I needed help with fieldwork or in the laboratory. I could not have done this without her. This research was supported in part by funding from the University of Guelph and the Natural Sciences and Engineering Research Council of Canada to J. D. A. TABLE OF CONTENTS General Introduction 1 Chapter 1: Biological and ecological mechanisms for overcoming sperm limitation in a freshwater mussel Abstract 22 Introduction 23 Methods 25 Results 32 Discussion 37 Chapter 2: The effect of near-bed turbulence on sperm dilution and fertilization success of broadcast spawning bivalves. Abstract 59 Introduction 60 Methods 62 Results 69 Discussion 76 Chapter 3: Effects of near-bed turbulence on settlement and resuspension of freshwater mussel larvae Abstract 99 Introduction 100 Methods 102 Results 108 u Discussion .1.14 Chapter 4: The role of bottom roughness parameters on the transport of sperm and larvae of benthic organisms Abstract 133 Introduction 134 Methods 136 Results 140 Discussion 143 General Conclusions 162 111 LIST OF TABLES Table II. Factors influencing fertilization success for broadcast spawners (from Okubo et al. 2001). 9 Table 3.1. Significant associations with R values greater than 0.100 for laboratory and field data using linear regression analysis. 124 LIST OF FIGURES Figure II. Forces acting on a static body. U represents direction of flow. 3 Figure 12. Vertical profile of a benthic boundary layer (from Nowell and Jumars 1984). 6 Figure 13. An example of an acoustic Doppler velocimeter velocity time series (taken 30 cm above the bed at Evans Point, Lake Erie on August 9, 2008) indicating an example of a fluctuation in the stream-wise velocity component, U. 6 Figure 14. Diagram of the u'w' quadrant plane (based on Cellino and Lemmin 2004). 6 Figure 15. Life history stages of Dreissena polymorpa (from Ackerman etal. 1994). : 13 Figure 1.1. (A) Results from the polyspermy study on dreissenid fertilization. Freshly spawned eggs and sperm were mixed and monitored over a 24 h period, with the proportion of successfully developing zygotes/embryos recorded at each time interval. (B) The stages of zygote development at; 0 h - unfertilized egg; 0.5 - fertilization envelope; 1.0 h - 2-cell stage; 2.0 h - 4-cell stage; 3.0 h - 8-cell stage; and at 24 h - trochophore larval stage. 50 Figure 1.2. Fertilization success as a function of sperm concentration for Dreissena polymorpha and Dreissena bugensis. Fertilization success increases relative to the increase in sperm concentration. Plots are means ± SE, N D. polymorpha = 8, N D.bugensts= 4. There was no significant difference between species. 51 Figure 1.3. Fertilization success as a function of egg concentration for Dreissena polymorpha and Dreissena bugensis. There was no significant relationship between fertilization success and egg concentration for either species. 52 Figure 1.4. Fertilization success in Dreissena polymorpha and Dreissena bugensis as a function of time post spawning for five sperm concentrations. Regardless of concentration, there is a decrease in fertilization success as sperm age. Values are means ± oil. IN D .polymorpha ~ O, FN o. bugensis ~ J- ?.J Figure 1.5. Results from COMSOL modeling for the mussel cluster scenario (A) and coral head scenario (B). Flow is left to right, with lines representing velocity streamlines and areas of recirculation are rotating in a clockwise direction. Areas of recirculation are indicated by the presence of vortices, examples of which can be seen behind the individual mussel and reef elements. Scalar (1 mol ml"1) which was released from the downstream slope of the first mussel cluster (C) or coral head (D) is retained in the recirculation zone or transported downstream. Note the different spatial scales for the two scenarios. White line in panel D represents an example of a transect used for scalar concentration measurements. 54 v Figure 1.6. The effect of modeled roughness element height (HRei = HR/Z) on the upstream retention vs. downstream transport of scalar to the next element (i.e., downstream of the second roughness element). The dotted line indicates the point at which 50% of released scalar switches from downstream transport to upstream retention, Note that the mussel cluster and coral head element fit the curve as well. Values are means ± SE. 55 Figure 1.7.
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