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Influence of Maternal Effects and Environmental Conditions on Growth and Survival of Atlantic Cod (Gadus Morhua L.)

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Influence of Maternal Effects and Environmental Conditions on Growth and Survival of Atlantic Cod (Gadus Morhua L.) Aus dem Leibniz Institut für Meereswissenschaften An der Christian-Albrechts-Universität zu Kiel Influence of maternal effects and environmental conditions on growth and survival of Atlantic cod (Gadus morhua L.) Dissertation zur Erlangung des Doktorgrades der Mathematisch-Naturwissenschaftlichen Fakultät der Christian Albrechts-Universität zu Kiel vorgelegt von Vivian Christiane Buehler Kiel 2004 Influence of maternal effects and environmental conditions on growth and survival of Atlantic cod (Gadus morhua L.) Table of Contents: Acknowledgements 1 1. Introduction 1 1.1 Importance of cod fisheries. 3 1.2 The Northeast Artic cod 4 1.2.1 Characteristics of the NEAC Stock 5 1.2.2 Sex ratio and spawning behaviour of NEAC 6 1.2.3 The offspring of NEAC 7 1.3 Factors that may influence stock management 8 1.4 What is maternal effect? 9 1.5 The MACOM Project 12 2. Materials and Methods 12 2.1 Parental Stock 12 2.1.1 Collecting Fish 12 2.1.2 Breeding 13 2.1.3 Incubation 14 2.1.4 Egg quality measurements 14 2.2 Mesocosm Experiment 14 2.2.1 Characteristics of the mesocosms 15 2.2.2 Monitoring of biotic and abiotic parameters 16 2.2.3 Transport and arrival of the larvae in Flødevigen 17 2.2.4 Releasing of larvae in the mesocosms 18 2.2.5 Sampling of cod larvae in the mesocosms 18 2.2.6 Termination of the Mesocosm Experiment 18 2.2.7 Mortality Calculation 19 2.3 Rearing of Juveniles in Indoor Tanks 20 2.3.1 Fish Tagging and fin clipping 20 2.3.2 Fish Measurements 21 2.3.3 Termination of the tanks Experiment 21 2.3.3.1 Estimation of oocytes size and Potential Fecundity 22 2.4 Laboratory Examinations 23 2.4.1 Otolith microstructure analysis 23 2.4.1.1 Otolith Preparation 23 2.4.1.2 Otolith reading 24 2.4.2 RNA/DNA ratio and glycolytic enzymes 24 2.4.3 DNA Fingerprinting 26 2.5 Statistical Analysis 27 3. Results 27 3.1 Description of the broodstock 29 3.2 Spawning and Egg quality parameters 35 3.3 Laboratory control 35 3.3.1 Mortality test on starving groups 35 3.3.2 Relations among egg size, larval size and condition 36 3.3.2.1 Newly-hatched otoliths 39 3.4 Mesocosm Experiment 39 3.4.1 Monitoring of the mesocosms 43 3.4.2 Larvae sampling and survival 53 3.4.3 Analysis of Growth and Condition 76 3.5 Tanks Period 84 4. Discussion 84 4.1 Exploitation and changes in demographic structure 85 4.1.2 Evolutionary Consequences of the shift in demographic structure 87 4.2 The influence of maternal effects 88 4.2.1 Detecting maternal effects in this study 89 4.2.1.1 Egg quality parameters 90 4.2.1.2 The offspring 90 4.2.1.3 Effect on otoliths 91 4.2.1.4 Biochemical methods 91 4.2.1.5 Tanks data 92 4.3 The connection between maternal effects and environment 92 4.4 Environmental effects 95 4.5 Analysis on the approach followed by this study 95 4.5.1 Differences between repeat and first spawners 95 4.5.2 Mating selection 96 4.5.3 Egg parameters 97 4.6 Evolutionary consequences of maternal effects 98 4.7 Conclusions 99 5. Summary 100 6. Zusammenfassung 101 7. References List of Figures List of Tables There is no end, no beginning. There is only the infinite passion of life… (Frederico Fellini) This work is dedicated to the maternal effects of all mothers that made of this world a better place to be… Acknowledgements I am very grate to Prof. Dr. Dietrich Schnack for the kindly supervision of this work, for his comprehension and patience; and to Prof. Dr. Erlend Moksness for giving me the opportunity to work in MACOM, thrusting me for so much responsibility in this project. Dr. Catriona Clemmesen, I thank for all the nice moments and fun we had in this project. This work could not be performed without the valuable contribution of all MACOM participants: Dr. Gary Carvalho, Dr. William Hutchinson, Dr. Lorenz Hauser and Richard Case were responsible for the genetic fingerprints. Dr. Olav Kjesbu, Dr. Håkon Otterå, Dr. Anders Thorsen were responsible for the broodstock and fecundity data. Dr. Catriona Clemmesen was responsible for all biochemical analyses. Dr. Helge Paulsen was responsible for the egg quality measurements and partially for the 10 week otoliths data. In addition, I would like to thank the coordinator of the project Dr. Terje Svåsand, Dr. Per Solemdal, and Dr. Tore Jakobsen. My best thanks to Jan Pedersen and Jon Kåre Stordal from Parisvatnet and all the staff from Flødevigen. Kjersti Eline Larsen, I thank for the nice teamwork in Norway. I specially thank Else Torstensen and Bente Lundin for friendship and support. Dr. Halvor Knutsen teached me the PCR and genetic fingerprinting methods. I learned a lot about Phytoplankton from Einar Dahl. Øystein Paulsen took many of the pictures in this project. Prof. Dr. Torfinn Lindem, Dr. Birgit Dannevig, Kitty and Christopher Ottersland, Christine Elmenhorst, Alf Dannevig, Nikolai Dannevig, Petter Baardsen, Tor Birkeland, Helen Nilsen and Inger Henriksen thanks for a great time in Norway. Dr. Odd Aksel Bergstad knows the most amazing stories about cod fisheries in Norway. Jan Pedersen, and the people from the University of Bergen helped me to identify a bacterial contamination and to control the outburst of vibriosis in juvenile cod during tank rearing. Prof. Dr. H. Rosenthal, Dr. Uwe Waller, Dr. Arild Folkvord and Dr. Ed trippel gave me great advices on aquaculture and fisheries. Dr. Hans-J. Krambeck, Herbert Kiesewetter and Werner Wegner from the Max-Planck-Institute for Limnology in Plön (MPIL), saved my work from many viruses and computer crashes. All my love to Prof. Dr. Wolfgang J. Junk, Prof. Dr. Ulrich Sommer and Prof. Dr. Joachim Adis for all the good advices, friendship and support. I thank Dr. Frank J. Jochem (Florida International University, US) for helping me out in many questions concerning phyto-zooplankton interactions and environmental factors. Prof. Dr. Joachim Gröger (University of Massachusetts, US) developed a nice model to test the complex MACOM data. Bene Willert and Helgi Mempel for the nice teamwork. Martin Lembke and Barbara Schmidt transformed every visit to the IfM library in a happy event. Daniel Stepputtis and Jörn Schmitt were a great help with software and graphic design. Bastian Huwer has been always a great friend and indispensable help during the final moths of this thesis. Brigitte Rohloff and her husband Kalle are the most phantastic people, who are able to charm a smile on your face even in the cloudiest days. Isabella Galvao was a nice company in the early hours at the IfM. Rudi Lüthje and Dr. Rudi Voss reminded me that there is always a “lighthouse” somewhere out there. From Muklis Kamal I’ve learned that friendship can sprout in the most different soils. Su-Kyoung Kim seasoned our lives with the most fragrant Asian flavors. Mary Stradas believe that everyone can be the alchemist of his own life... Finally, I thank my family, friends and colleagues for their comprehension, love and support. Introduction__________________________________________________________________ 1. Introduction Gadus morhua, the codfish (fig.1) belongs to the family Gadidae, order Gadiformes, class Actinopterygii (ray-finned fishes). The maximum historical size reported was 200 cm of total length, 96 Kg, and 25 years of age (Cohen et al. 1990). Cod live in a benthopelagic, oceanodromous, marine or brackish environment with depths ranging from 1 to 600 m. The geographic distribution of the species is the North Atlantic ranging from 35° to 78° N of latitude, i.e. from Cape Hatteras to Ungava Bay along the North American coast, east and west of Greenland, around Iceland; coasts of Europe from the Bay of Biscay to the Barents Sea. Additionally, two stocks are recognized in the brackish Baltic Sea. The distinguishing features are a stout body with a long chin’s barbel, relatively small eyes, and three dorsal fins all close together at the base and rounded in outline, and two anal fins. The upper jaw overhangs the lower. The lateral line is continuous, with a smooth curve about the pectoral fin. The color varies according to the local habitat and ranges from light to brown leopard spots on an olive to gray- greenish or sometimes even reddish back. They are white ventrally and the lateral line is conspicuous light. Cod are omnivorous and feeds at dusk and at dawn on invertebrates and fish, including young cod. Figure 1. Gadus morhua L. Cod are prized for their white flesh, which has only 0.3% fat and 18% protein per gram. Besides, there is almost no waste to a cod. The head is cooked in soups, while the throat (known as cod tongue) and cheeks are very appreciated in Norway. Additionally, gonads, stomachs, and livers are all eaten or used in many other products. The liver oil is highly valued for its vitamins D and E. 1.1 Importance of cod fisheries The story of cod fisheries looses itself in the history of times. Cod have been already called “Beef of the sea” and their history goes further back to the first registers during Viking times, or probably even earlier. Through the centuries, cod fisheries sustained different societies in both sides of the North Atlantic, and fisheries became more intense every year. The nineteenth century was marked by the optimism that natural resources would never end up. Although many fishermen started to complain about decreases in catches, Thomas Henry Huxley (1825-1895), a respected British biologist (also known as “Darwin’s Bulldog” for his defense on the theory of 1 Introduction__________________________________________________________________ evolution) argued in 1883 at the International Fisheries Exhibition in London that “overfishing” was an unscientific and erroneous fear.
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