FACTORS INFLUENCING ADULT GONAD PRODUCTION and LARVAL GROWTH and SURVIVAL of the PURPLE SEA URCHIN (Strongylocentrotus Purpuratu
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FACTORS INFLUENCING ADULT GONAD PRODUCTION AND LARVAL GROWTH AND SURVIVAL OF THE PURPLE SEA URCHIN (Strongylocentrotus purpuratus) by Md. Abul Kalam Azad M. Sc. (Fisheries Biology), Bangladesh Agricultural University, Bangladesh, 1997 M. Sc. (Aquaculture), Asian Institute of Technology, Thailand, 2002 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in The Faculty of Graduate Studies (Animal Science) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) August 2011 © Md. Abul Kalam Azad, 2011 Abstract The continued demand for sea-urchin gonads and overexploitation of natural stocks in many countries have stimulated interest in the aquaculture of various sea-urchin species. The overall aim of this thesis was to develop rearing protocols for larval and adult purple sea urchins (Strongylocentrotus purpuratus). The specific objectives of this thesis were to determine: (1) the effect of temperature and diet on adult gonad production and (2) the effect of temperature, microalgal diet, ration, and stocking density on embryonic/larval growth and survival. To test these objectives I measured ingestion rate, gravimetric absorption, assimilation efficiency, and various gonad attributes (i.e. wet weight, water content, gonad index, texture, firmness, colour, flavour, and maturity stage) in the experiments using adult sea urchins and developmental progression from egg to prism stage, embryo length, larval length, larval mid-line body length, larval body width, larval post-oral arm length, time to reach metamorphic competency, and survival rate in the embryo/larval experiments. Adult purple sea urchins produced the highest quality gonads at 12°C when fed a prepared diet developed specifically for urchin gonad production by the Norwegian Institute of Fisheries and Aquaculture. Embryos and larvae showed more normal development and had significantly higher percent survival when reared at 11 and 14°C than at 8 and 17°C with best growth and survival occurring at stocking densities ≤100 ind ml-1 of eggs/embryos and ≤1.0 ind ml-1 of larvae. The single-species algal diet of Dunaliella tertiolecta and the bi-species diet of D. tertiolecta and Isochrysis sp. supported the highest larval growth and percent survival of all phytoplankton diets evaluated. Larvae fed an increasing diet ration (i.e. 2,500–10,000 cells ml-1, according to developmental stage) of a mixed-species algal diet (D. tertiolecta and Isochrysis sp.) generally had significantly better growth and survival than any fixed rations (i.e. same ration throughout development) evaluated. Overall, the results from ii this study suggest that S. purpuratus could be an excellent potential candidate species for aquaculture development. iii Preface For this thesis, I contributed to the development of the experimental protocols, conducted the research, performed the statistical analyses, interpreted the data, and prepared the various chapters. I have written all parts of the manuscripts mentioned in this dissertation. All of my committee members (Dr. Abayomi Alabi, Dr. Colin Brauner, Dr. Marina von Keyserlingk, Dr. Christopher M. Pearce, and Dr. Robert Scott McKinley) helped to conceive the design of the original experiments. Drs. McKinley and Pearce provided overall supervision and guidance to me and ensured that the experiments were conducted appropriately and that data analyses were correctly performed. As co-authors, Drs. McKinley and Pearce reviewed and provided critical comments on all of the manuscripts included in this dissertation. A version of Chapter 1 has been published [Azad A.K., McKinley S. and Pearce C.M. (2010) Factors influencing the growth and survival of larval and juvenile echinoids. Reviews in Aquaculture 2: 121-137]. A version of Chapter 2 has been published [Azad A.K., Pearce C.M. and McKinley R.S. (2011) Effects of diet and temperature on ingestion, absorption, assimilation, gonad yield, and gonad quality of the purple sea urchin (Strongylocentrotus purpuratus). Aquaculture 317: 187-196]. A version of Chapters 3 and 4 has been published [Azad A.K., Pearce C.M. and McKinley R.S. (2011) Influences of stocking density and temperature on early development and survival of the purple sea urchin, Strongylocentrotus purpuratus (Stimpson, 1857). Aquaculture Research: in press]. A version of Chapters 5 and 6 has been accepted for publication [Azad A.K., Pearce C.M. and McKinley R.S. (2011) Influence of algal species and dietary rations on larval development and survival of the purple sea urchin, Strongylocentrotus purpuratus (Stimpson, 1857). Aquaculture: accepted]. iv Drs. Alabi, Brauner, and Keyserlingk provided critical comments on the manuscript in Chapter 1 as well as all of the chapters in this dissertation. Dr. Pearce secured introduction and transfer permits from the Pacific Region Introductions and Transfers Committee for the collection of experimental animals and obtained approval of protocols (Animal Use Protocol references : 08-012 and 09-020) for experimental use of animals from the Fisheries and Oceans Canada (DFO) Pacific Region Animal Care Committee (PRACC). Dr. Pearce provided laboratory space at the Pacific Biological Station and ensured that the experiments were conducted in accordance with the guidelines of the DFO PRACC and the guidelines of the Canadian Council on Animal Care (CCAC). v Table of contents Abstract ........................................................................................................................................... ii Preface.............................................................................................................................................iv Table of contents .............................................................................................................................vі List of tables.....................................................................................................................................x List of figures ................................................................................................................................ xii Acknowledgements .......................................................................................................................xvi Dedication .................................................................................................................................. xviii CHAPTER 1: Introduction ...........................................................................................................1 1.1 General introduction...............................................................................................................1 1.2 General biology and ecology of sea urchins ..........................................................................6 1.3 Aquaculture potential of sea urchins....................................................................................11 1.4 Factors influencing the growth and survival of larval and juvenile echinoids: a review.....13 1.4.1 Larval production ..........................................................................................................14 1.4.1.1 Food: phytoplankton ..............................................................................................14 1.4.1.2 Food: prepared diets...............................................................................................22 1.4.1.3 Food: dissolved organic matter ..............................................................................24 1.4.1.4 Stocking density .....................................................................................................25 1.4.1.5 Temperature ...........................................................................................................26 1.4.2 Juvenile production .......................................................................................................29 1.4.2.1 Food: algae .............................................................................................................29 1.4.2.2 Food: prepared diets...............................................................................................35 1.4.2.3 Stocking density .....................................................................................................38 1.4.2.4 Temperature ...........................................................................................................40 1.5 The experimental animal......................................................................................................42 1.6 Hypotheses and objectives ...................................................................................................45 vi 1.6.1 Experiment 1: Influence of diet and temperature on adult gonad production...............45 1.6.2 Experiment 2: Influence of stocking density on early development and survival ........46 1.6.3 Experiment 3: Influence of temperature on early development and survival ...............47 1.6.4 Experiment 4: Influence of algal species on larval growth and survival ......................47 1.6.5 Experiment 5: Influence of dietary rations on larval growth and survival....................48 CHAPTER 2: Influence of diet and temperature on gonad production of the purple sea urchin.............................................................................................................................................50 2.1 Introduction ..........................................................................................................................52