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Biology of European Sea Bass TThhiiss ppaaggee iinntteennttiioonnaallllyy lleefftt bbllaannkk Biology of European Sea Bass Editors F. Javier Sánchez Vázquez Department of Physiology University of Murcia Murcia Spain José A. Muñoz-Cueto Department of Biology University of Cadiz Cadiz Spain p, A SCIENCE PUBLISHERS BOOK GL--Prelims with new title page.indd ii 4/25/2012 9:52:40 AM CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2015 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Version Date: 20140602 International Standard Book Number-13: 978-1-4665-9946-8 (eBook - PDF) This book contains information obtained from authentic and highly regarded sources. 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For permission to photocopy or use material electronically from this work, please access www.copyright.com (http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com Preface European sea bass, Dicentrachus labrax, has attracted the attention of many people: scientists, fi sh farmers, fi shermen and other stakeholders. Since ancient times, this species has been regarded as a highly valuable table fi sh, and it is considered a delicacy in restaurants today. This fi sh has many different names. In France, it is called “loup de mer”, while in the Iberian Peninsula it is known as “lubina” or “robalo”. In Italy sea bass is called “branzino” or “spigola”, whereas the Greeks refer to this fi sh as “lavraki” (from its vernacular name) and the Turks as “levrek”. European sea bass was historically cultured in coastal lagoons and tidal reservoirs, so it is not surprising that this fi sh species was one of the fi rst marine fi sh to be farmed in Europe using modern aquaculture techniques. In fact, in Italy sea bass and seabream are the most consumed fi sh species. With declining fi sheries, the aquaculture of European sea bass becomes increasingly important to provide a high quality product to consumers. In fact, according to FAO predictions, by 2030 over 65% of seafood will be supplied by aquaculture. Sea bass aquaculture production has been rising since the early 90s, reaching over 130,000 tonnes per year in the last few years, while fi sheries catches remain stackled around 10,000 tonnes per year. Geographically, the main production of sea bass is located in two countries: Turkey (34.2%) and Greece (34.1%); although this species is also cultured in many other Mediterranean countries, such as Spain (11.4%), Italy (6.9%), Egypt (4%), Croatia (2.5%), France (2.4%), Tunisia, Cyprus, Portugal, etc. In the last few decades, many researchers devoted their investigations to this species, publishing over 2,200 papers in scientifi c journals. Most of this research was conducted in three European countries: France (466), Spain (384) or Italy (361); followed by Portugal (136), Greece (133) and Turkey (99). The most popular research areas are Fisheries (including Aquaculture), Zoology and Marine & Freshwater Biology, with many signifi cant contributions in the specifi c fi elds of (in relevance order) Physiology, Genetics, Immunology, Ecology, Endocrinology, etc. In this book, we have focused on the general Biology, Ecology, Physiology, Behavior and Pathology of the European sea bass. The book is organized in twelve chapters authored by some of the world’s most renowned scientists in these fi elds. vi Biology of European Sea Bass The book starts with a chapter on the “Ecology and Distribution of Dicentrarchus labrax (Linnaeus 1758)”, written by Ángel Pérez-Ruzafa and Concepción Marcos, from the Department of Ecology and Hydrology, University of Murcia (Spain). In this chapter, the authors provide information on the distribution and genetic structure of sea bass, the main environmental factors affecting sea bass distribution and behavior and the synergetic effect of multiple factors on physiological processes, which determines the way in which this species copes with environmental variability. Other interesting aspects such as migration, reproduction, fecundity and reproductive efforts, recruitment, feeding ecology and behavior, life history strategies, response to pollution, management and conservation of sea bass are also addressed in this chapter. Most living organisms have developed a biological clock to keep time and cope with predictable environmental changes by anticipating the forthcoming of a recurrent event in their habitat. Fish in general, and sea bass in particular, are no exception to this rule. The second chapter entitled “The Biological Clock and Dualism” examines this topic; it has been authored by Ana del Pozo and F. Javier Sánchez-Vázquez, from the Department of Physiology, University of Murcia (Spain) and Jack Falcón, from the Banyuls-sur-Mer Oceanological Observatory, CNRS-UPMC (France). This chapter focuses on biological rhythms and zeitgebers (food, photoperiod, temperature), the molecular clock machinery that controls these rhythms, as well as in dualism (diurnal vs. nocturnal) and the mechanisms controlling this dualism in feeding and locomotor behavior, digestive physiology, and seasonal reproductive rhythms in sea bass. The pineal organ of fish is a photosensory and neuroendocrine epithalamic structure that plays a key role in the temporal organisation of physiological and behavioral processes. In the chapter “The Pineal Organ of the European Sea Bass: A Neuroanatomical Approach”, the authors present a review of their studies on the macroscopic and microscopic anatomy of the pineal organ in the European sea bass and the characterization of pineal photoreceptive and melatonin-synthesizing cells. Moreover, they summarise the bidirectional connections of the pineal with the brain through pinealofugal (efferent) and pinealopetal (afferent) projections, as well as the targets of the neuroendocrine message of the pineal organ (melatonin) by describing the tissue-specifi c distribution and localization melatonin receptors in the retina, the brain and somatic tissues of sea bass. This chapter has been written by Patricia Herrera-Pérez, M. Carmen Rendón- Unceta and José Antonio Muñoz-Cueto, from the Department of Biology of the University of Cadiz (Spain), Arianna Servili, from the Department of Functional Physiology of Marine Organisms, IFREMER (France) and Jack Falcón, from the Banyuls-sur-Mer Oceanological Observatory, CNRS- UPMC (France). Preface vii The following chapter is entitled “Melatonin Rhythms” and is authored by Luisa M. Vera and Hervé Migaud, from the Institute of Aquaculture, University of Stirling (UK). In the European sea bass, several studies have focused on the description, regulation and physiological role of melatonin rhythms. Current knowledge covers issues regarding photoreception, infl uence of light intensity and spectrum on melatonin production, circadian and seasonal rhythmicity, extrapineal sources of melatonin, distribution, and daily and seasonal rhythms of melatonin receptors, among others. Many of these fi ndings are reviewed and discussed throughout this chapter. Fish integrate environmental cues using specifi c sensory systems such as the pineal organ, and transduce them into a cascade of hormones along the brain-pituitary-gonadal axis, which control the timing of seasonal reproduction. The purpose of this sequence of events is to ensure that reproduction takes place at the time of year most favourable for the survival of progeny. In the next chapter, entitled “Neuroendocrine Regulation of Reproduction in Sea Bass (Dicentrarchus Labrax)”, the authors focus on the extraordinary complexity of the neuroendocrine control of reproductive axis of sea bass and they detail the factors modulating this reproductive axis, with both positive and negative effects, including the relevant knowledge acquired very recently in their laboratories. This chapter is authored by Arianna Servili, from the Department of Functional Physiology of Marine Organisms, IFREMER (France), Olivier Kah, from INSERM, Université de Rennes 1-CNRS (France) and Patricia Herrera-Pérez, José Antonio Paullada Salmerón and José Antonio Muñoz-Cueto, from the Department of Biology of the University of Cadiz (Spain). During the fi rst weeks of life, marine fi sh larvae undergo signifi cant morphological and physiological modifi cations to acquire all the adult features by the end of the larval period. This phase is certainly the most critical in the life cycle of fi sh, not only in natural environments, but also under controlled aquaculture conditions. The chapter “European Sea Bass Larval Culture” represents the contribution of Enric Gisbert and Alicia Estévez, from IRTA Centre de Sant Carles de la Ràpita (Spain), Ignacio Fernández, from CCMAR/CIMAR, Universidade do Algarve (Portugal), Natalia Villamizar, from the Department of Physiology, University of Murcia (Spain), Maria J. Darias, from IRD-Université Montpellier II (France) and Jose L. Zambonino-Infante, from the Department of Functional Physiology of Marine Organisms, IFREMER (France). To begin, the authors focus on the embryonic and larval development of European sea bass with a comprehensive analysis of the ontogenetic changes during the early life stages of fi sh. Later, a description of the current larval production systems at an industrial and laboratory scale is presented with special emphasis on the optimal rearing factors affecting fry production. Finally, the authors viii Biology of European Sea Bass review how some of the most important abiotic (light and temperature) and biotic (nutrition) factors affect larval performance. Feeding by fi sh is dependent upon their sensory capacities to locate food, their ability to capture, handle and ingest food items, and their physiological and biochemical capacities to digest and transform ingested nutrients. In the chapter “Foraging Behavior”, Sandie Millot and Marie- Laure Begout from the Fishing Resources Laboratory of La Rochelle, IFREMER (France) and David Benhaïm, from LERMA, INTECHMER/ CNAM, Cherbourg (France), describe the main traits of foraging behavior both in the wild and in aquaculture for the different life stages of sea bass. The chapter also presents foraging behavior in the natural environment, then details foraging in the aquaculture environment in relation with environmental factors and fi nally describes other related behavior and illustrates the infl uence of domestication and selection processes on sea bass behavioral traits. Feeding is a key function in evolution since metabolic energy obtainment and storage make an important contribution to survival rates. The following chapter entitled “Food Intake Regulation”, is aimed at reviewing all the central and peripheral regulators of food intake that have been studied in sea bass. In addition, the author also reviews the relationship of food intake with some physiological processes (such as reproduction and stress response) and environmental factors (temperature, salinity, oxygen and ammonia) which are known to modulate food intake in sea bass. The author of this chapter is José Miguel Cerdá-Reverter, from the Institute of Aquaculture of Torre de la Sal, CSIC (Spain). In fi sh farming, the cost of feed is the largest factor of production, and, therefore, it is important to minimize the cost of the feed while maintaining high growth rate of the fi sh. In this context, the chapter “Nutrition and Dietary Selection” represents the contribution of Rodrigo Fortes da Silva, Francisco Javier Sánchez-Vázquez and Francisco Javier Martínez-López, from the Department of Physiology of the University of Murcia (Spain). The fi rst section of this chapter deals with basic and practical aspects, comprising protein, energy, lipid and carbohydrate nutrition in sea bass. The requirements of each macronutrient and its replacement with sustainable food sources, micronutrient requirements and the use of probiotics are also reviewed. The second section of the chapter covers dietary selection and the ability of sea bass to detect and self-select different food types. Both macro- and micro-nutrient selection are considered, as well the basic mechanisms responsible for the control of food intake and dietary selection in sea bass. Farming practices alter the natural equilibrium between the host and the pathogen found in the wild, favoring the emergence of diseases and posing a major problem for the aquaculture industry. Farmed fi sh are also Preface ix constantly exposed to different types of stressors such as bad or poor culture conditions, inadequate diets and other anthropogenic factors that might compromise their performance and survival. In the chapter “Pathology”, Ariadna Sitjà-Bobadilla, from the Institute of Aquaculture of Torre de la Sal, CSIC (Spain), Carlos Zarza, from Tassal Group Limited, Tasmania (Australia) and Belén Fouz, from the Department of Microbiology and Ecology, University of Valencia (Spain), review the main pathologies due to bacteria, virus, protozoan and metazoan parasites in the European sea bass. In addition, they also discuss other pathologies induced by aquaculture practices, disorders provoked by harmful algae blooms and jellyfi sh and tumors reported in sea bass populations. The fish immune system is characterized by a multi-layered organization that provides immunity to infectious organisms, and each layer can be considered to have an increasing complexity. In this way the central challenge of fi sh immunology, which includes the sea bass model, is now to fully characterize the occurring immune responses. The next chapter is entitled “Current Knowledge on the Development and Functionality of Immune Responses in the European Sea Bass (Dicentrarchus labrax)” and is authored by Jorge Galindo-Villegas and Victoriano Mulero, from the Department of Cell Biology and Histology, University of Murcia (Spain). In this chapter, the authors briefl y summarize and update the emergence, phylogenetic distribution and functionality of D. labrax innate and adaptive immunity, as well as the most relevant interactions between them towards host defense against pathogens. The book ends with a chapter on “The Response to Stressors in the Sea Bass”. Stress is an important physiological response that all animals, including fi sh, experience when they are subjected to alarming situations. The stress impact in fi sh has been assessed in aquaculture in relation to fi sh transport, environmental factors, and factors related to husbandry practices, including social stressors. The issue of stress in aquaculture is highly signifi cant, because a number of consequences often undesired but always relevant for fi sh performance such as growth, disease resistance, behavior and energy balance are clearly observed. Many of the fi ndings obtained by authors in relation to the stress response in the European sea bass are reviewed and discussed in this chapter. The authors contributing to this chapter are Lluis Tort, from the Department of Cell Biology, Physiology and Immunology, Autonomous University of Barcelona, (Spain), Josep Rotllant, from the Vigo Marine Research Institute, CSIC (Spain), Michalis Pavlidis, from the Department of Biology, University of Crete (Greece), Daniel Montero, from the Aquaculture Research Group, University of Las Palmas de Gran Canaria (Spain) and Genciana Terova, from the Department of Biotechnology and Life Sciences, University of Insubria (Italy).