Biogeochemistry of Marine Systems (Sheffield Biological Sciences S.)

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Biogeochemistry of Marine Systems

Biological Sciences Series A series which provides an accessible source of information at research and professional level in chosen sectors of the biological sciences. Series Editors: Professor Jeremy A. Roberts, Plant Science Division, School of Biosciences, University of Nottingham. Professor Peter N.R. Usherwood, Molecular Toxicology Division, School of Biosciences, University of Nottingham. Titles in the series: Stress Physiology in Animals Edited by P.H.M. Balm Seed Technology and its Biological Basis Edited by M. Black and J.D. Bewley Leaf Development and Canopy Growth Edited by B. Marshall and J.A. Roberts Environmental Impacts of Aquaculture Edited by K.D. Black Herbicides and their Mechanisms of Action Edited by A.H. Cobb and R.C. Kirkwood The Plant Cell Cycle and its Interfaces Edited by D. Francis Meristematic Tissues in Plant Growth and Development Edited by M.T. McManus and B.E. Veit Fruit Quality and its Biological Basis Edited by M. Knee Pectins and their Manipulation Edited by G.B. Seymour and J.P. Knox Biogeochemistry of Marine Systems Edited by K.D. Black and G.B. Shimmield

Biogeochemistry of Marine Systems Edited by KENNETH D. BLACK Scottish Association for Marine Science Dunstaffnage Marine Laboratory Oban, UK and GRAHAM B. SHIMMIELD Director, Scottish Association for Marine Science Dunstaffnage Marine Laboratory Oban, UK

Blackwell Publishing

© 2003 by Blackwell Publishing Ltd Editorial Offices: Blackwell Publishing Ltd, 9600 Garsington Road, Oxford OX4 2DQ, UK Tel: +44 (0)1865 776868 Blackwell Publishing Asia Pty Ltd, 550 Swanston Street, Carlton, Victoria 3053, Australia Tel: +61 (0)3 8359 1011 ISBN 1–84127–327–9 Originated as Sheffield Academic Press Published in the USA and Canada (only) by CRC Press LLC, 2000 Corporate Blvd., N.W., Boca Raton, FL 33431, USA Orders from the USA and Canada (only) to CRC Press LLC USA and Canada only: ISBN 0–8493–2818–7 The right of the Author to be identified as the Author of this Work has been asserted in accordance with the Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use.

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Contents

Preface Contributors 1 Mangroves of Southeast Asia

xii xv 1

M. HOLMER 1.1 1.2 1.3 1.4

Introduction Mangrove forest structure and function Water column biogeochemistry Organic matter sources in mangrove forests 1.4.1 Decomposition of detritus 1.5 Sediment biogeochemistry 1.5.1 Total microbial activity in mangrove sediments 1.5.2 Mineralization pathways in mangrove sediments 1.5.3 Phosphorus cycling 1.6 Factors influencing the biogeochemistry 1.6.1 Effect of forest type and age 1.6.2 Influence of macrofauna 1.6.3 Effect of seasonal variations on mangrove forest biogeochemistry 1.7 Sediment biogeochemistry and implications for mangrove vegetation 1.8 Biogeochemistry in mangroves affected by anthropogenic activities References 2 Coral reefs

1 2 3 7 9 11 12 15 23 25 25 26 28 29 31 34 40

M.J. ATKINSON and J.L. FALTER 2.1 Introduction 2.2 Coral reef morphology and zonation 2.3 Basic biogeochemistry 2.3.1 Carbon 2.3.2 Dissolved organic matter 2.3.3 Nitrogen 2.3.4 Phosphorus

40 41 43 43 47 48 50

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CONTENTS

2.3.5 Silica 2.3.6 Iodine 2.4 Interstitial geochemistry and hydrology of coral reef frameworks 2.5 Mass transfer-limited biogeochemical rates 2.6 Coral growth in high nutrient water 2.7 Measurement techniques 2.8 Summary statements References 3 Fjords

51 51 52 54 56 58 59 59 65

J.M. SKEI, B. MCKEE and B. SUNDBY 3.1 Introduction 3.1.1 Definition and origin of fjords 3.1.2 The public and scientific interest in fjords 3.2 Sediment diagenesis in oxic fjords 3.2.1 The Saguenay Fjord 3.2.2 Sedimentation 3.2.3 Composition of the rapidly deposited layers 3.2.4 Sulfate reduction and sulfide accumulation 3.2.5 Mercury diagenesis 3.2.6 Phosphorus and arsenic geochemistry 3.2.7 Non-steady-state diagenesis 3.3 Elemental cycling in anoxic waters 3.3.1 Chemical tracers 3.3.2 Cycling of carbon and nutrients 3.3.3 Trace element and radionuclide cycling 3.3.4 Fe–S systematics 3.3.5 Sulfate reduction and methane oxidation 3.3.6 Elemental cycling in sediments underlying anoxic waters 3.3.7 Preservation of organic matter References 4 The Eastern Mediterranean

65 66 67 69 69 69 70 70 73 73 75 76 76 77 78 81 83 84 85 86 91

MICHAEL D. KROM, STEVEN GROOM and TAMAR ZOHARY 4.1 Introduction 4.2 History of the Mediterranean basin 4.3 Basic description of the bathymetry and physical oceanography of the Eastern Mediterranean 4.3.1 Bathymetry

91 92 93 93

CONTENTS

4.3.2 Physical circulation of the Eastern Mediterranean 4.3.2.1 Formation of LDW 4.3.2.2 Formation of LIW 4.3.3 Recent water mass changes in the Eastern Mediterranean 4.3.4 Current patterns 4.4 Nutrients and chlorophyll distribution across the Eastern Mediterranean 4.4.1 General comments 4.4.2 Seasonal distributions 4.4.2.1 Winter 4.4.2.2 Spring into summer 4.4.3 Nutrient distribution below the nutricline 4.5 Total chlorophyll distribution and characteristics 4.5.1 Light penetration 4.5.2 Species composition 4.5.2.1 The prochlorophytes 4.5.2.2 The unicellular cyanobacteria 4.5.2.3 The eukaryotes 4.5.2.4 Heterotrophic bacteria 4.6 Primary production 4.6.1 Gradient in biomass and productivity from coastal waters to the open sea 4.7 Effects of mesoscale features on nutrient and chlorophyll distribution and phytoplankton productivity 4.7.1 Biogeochemical processes in mesoscale features 4.7.1.1 Rhodes cold-core (cyclonic) eddy 4.7.1.2 Cyprus warm-core (anti-cyclonic) eddy 4.7.1.3 Effects of other mesoscale features 4.8 Seasonal changes in phytoplankton biomass as detected by remote sensing 4.9 Nutrient limitation in the Eastern Mediterranean 4.10 Magnitude of man-induced changes in nutrient inputs and their possible effects on the Eastern Mediterranean 4.11 Summary and conclusions Acknowledgements Glossary References 5 The Arctic seas

vii 93 94 95 96 97 98 98 98 98 100 100 100 102 102 102 103 104 104 105 107 108 108 108 112 113 113 116 118 120 121 122 122 127

MICHAEL L. CARROLL and JOLYNN CARROLL 5.1 5.2

Summary Main features

127 128

viii

CONTENTS

5.2.1 Water masses 5.2.2 Continental shelves 5.2.3 Sea ice 5.3 Biogeochemical cycles and ecological processes 5.4 Environmental changes 5.4.1 Climate variability 5.4.2 Long-term climate change 5.4.3 Ozone and ultraviolet radiation 5.4.4 Contaminants 5.5 Natural resources and ecological services 5.5.1 Indigenous people 5.5.2 Non-indigenous regional populations 5.5.3 National/international/global users Acknowledgements References 6 The Arabian Sea

128 131 132 133 138 138 140 143 143 144 144 145 146 147 147 157

S.W.A. NAQVI, HEMA NAIK and P.V. NARVEKAR 6.1 6.2 6.3 6.4

Introduction Geographical setting Climate and circulation Nutrients and primary production 6.4.1 Subsurface nutrient trap 6.4.2 Primary productivity 6.4.3 New production 6.4.4 Phytoplankton composition and size distribution 6.4.5 Chlorophyll and POC 6.4.6 Effect of changes in mixed layer depth 6.5 Heterotrophic biomass and production 6.5.1 Heterotrophic bacteria 6.5.2 Nano- and microheterotrophs 6.5.3 Mesozooplankton 6.6 Food web structure and export of material to the deep sea 6.6.1 Phytoplankton growth and mortality 6.6.2 Particle fluxes to deep sea 6.6.3 Role of Arabian Sea as a source or sink of carbon dioxide (CO2) 6.7 Oxygen-deficient zones 6.7.1 Denitrification 6.7.2 Intermediate nepheloid layer 6.7.3 Other redox-sensitive elements 6.7.4 Biological effects

157 157 158 163 163 164 166 167 171 172 174 174 175 177 179 179 180 185 185 186 191 192 193

CONTENTS

6.8 Benthic processes References 7 The northeastern Pacific abyssal plain

ix 195 198 208

ANGELOS K. HANNIDES and CRAIG R. SMITH 7.1 Introduction 7.2 Key habitat parameters of deep seafloor communities 7.2.1 Key habitat parameters 7.2.1.1 Substratum type 7.2.1.2 Near-bottom currents 7.2.1.3 Bottom-water oxygen 7.2.1.4 Sinking POC flux 7.2.1.5 Redox conditions 7.2.2 Variation of key habitat parameters in the northeastern Pacific abyssal plain 7.2.2.1 Sediment types 7.2.2.2 Near-bottom currents and oxygen concentrations 7.2.2.3 POC flux and redox conditions 7.3 Northeastern Pacific abyssal zones 7.3.1 The eutrophic equatorial abyss 7.3.2 The mesotrophic (sub-equatorial) abyss 7.3.3 The oligotrophic central gyre abyss 7.4 Sensitivity and resilience to natural and anthropogenic change 7.4.1 General thoughts 7.4.2 Potential sensitivity and resilience to specific changes 7.4.2.1 Climate variation in the equatorial and North Pacific 7.4.2.2 Global increase in atmospheric greenhouse gases and temperatures 7.4.2.3 Manganese nodule mining 7.4.2.4 Iron fertilization 7.5 Concluding remarks Acknowledgments References 8 Deep-sea hydrothermal vents and cold seeps

208 208 209 209 210 211 211 211 212 212 212 213 213 214 217 218 220 220 221 221 223 226 229 230 231 231 238

RICHARD J. LÉVEILLÉ and S. KIM JUNIPER 8.1 Introduction 8.1.1 Deep-sea hydrothermal vents and cold seeps 8.1.2 Life at hydrothermal vents and cold seeps 8.1.3 Scope of this chapter

238 238 238 240

x

CONTENTS

8.2 Deep-sea hydrothermal vents 8.2.1 Distribution and general characteristics 8.2.1.1 Geochemical fluxes of gases and elements from hydrothermal vents 8.2.1.2 Off-axis diffuse flow versus axial venting 8.2.2 Subsurface biosphere at mid-ocean ridges 8.2.2.1 Evidence for a subsurface biosphere at deep-sea hydrothermal vents 8.2.2.2 Biogeochemical interactions in subsurface environments 8.2.3 Seafloor microbe-mineral interactions at hydrothermal vents 8.2.3.1 Microbial distribution and activity at vents 8.2.3.2 Biomineralisation at vents 8.2.3.3 Fossilisation of microbes at vents 8.2.3.4 Bacterial weathering of sulphides 8.2.4 Biogeochemical interactions in hydrothermal plumes 8.2.4.1 General features of hydrothermal plumes 8.2.4.2 Microbial ecology of hydrothermal plumes 8.2.4.3 Microbial productivity and organic carbon in plumes 8.2.4.4 Biogeochemical interactions in plumes 8.2.5 Biogeochemistry of off-axis vents and seafloor basalt 8.2.5.1 Off-axis vents 8.2.5.2 Seafloor basalts 8.3 Deep-sea cold seeps 8.3.1 Distribution, occurrences and general characteristics 8.3.1.1 Gas hydrates 8.3.1.2 Geochemical fluxes 8.3.2 Biogeochemistry of seep sediment pore fluids 8.3.2.1 Methanogenesis 8.3.2.2 Anaerobic sulphate reduction 8.3.2.3 Aerobic microbial oxidation of sulphide and methane 8.3.2.4 Anaerobic oxidation of methane 8.3.3 Microbial carbonates 8.4 Stability and perturbations of seafloor hydrothermal vent and cold seep systems 8.4.1 Geological stability of vents and seeps 8.4.2 Future perturbations related to resource extraction 8.4.2.1 Hydrothermal sulphides 8.4.2.2 Cold seeps 8.4.3 Response of cold seeps and gas hydrates to global warming

241 241 243 245 246 246 251 252 252 256 260 260 261 261 262 263 264 265 265 266 267 267 268 269 270 271 271 272 273 274 276 276 277 277 278 278

CONTENTS

8.5 Future work 8.6 Conclusion References 9 Influence of nutrient biogeochemistry on the ecology of northwest European shelf seas

xi 279 282 282

293

PAUL TETT, DAVID HYDES and RICHARD SANDERS 9.1 Introduction 9.2 Nutrient cycles 9.2.1 Macronutrient element availability 9.2.2 Sources of macronutrients 9.2.3 Sinks of macronutrients 9.2.4 Observed distributions of macronutrient concentrations and ratios 9.2.5 Iron 9.3 Plankton biogeochemistry 9.3.1 Taxonomy and life forms in the plankton 9.3.2 Theories of floristic composition 9.3.2.1 Light-nutrient-mixing explanations 9.3.2.2 Biogeochemical controls 9.3.2.3 Ecological controls 9.3.3 Variation in nutrient element ratios and its explanation in terms of plankton biochemistry 9.3.4 Quantitative theory for nutrient element ratios 9.3.5 Differences in abilities to assimilate different nutrients 9.3.6 Theoretical conclusions 9.4 Effects of ambient nutrient ratios on plankton 9.4.1 Introduction 9.4.2 Time series: Helgoland and the German Bight 9.4.3 Mesocosm and other competition experiments 9.4.4 Observations at sea 9.5 Discussion and conclusions 9.5.1 Introduction 9.5.2 Do high ambient N:Si ratios favour flagellates? 9.5.3 Do non-Redfield ambient N:P ratios perturb pelagic ecosystems? 9.5.4 The possibility of iron limitation in shelf seas 9.5.5 Trophic consequences of ratio changes – a Panglossian conclusion? 9.5.6 A flexible Redfield ratio? Dedication References Index

293 294 294 296 298 300 301 303 303 308 309 310 312 315 320 325 327 327 327 329 332 337 341 341 342 344 345 345 347 350 351 364

Preface

Marine biogeochemistry is a broad, interdisciplinary subject overlapping a range of other disciplines such as marine chemistry, geochemistry, ecology, physiology and oceanography, but in its own right it has become pivotal to progress in marine research in recent years. As a key component of the ‘earth system’, marine biogeochemistry interfaces directly with terrestrial, atmospheric and geological sciences. A working definition of the subject might be ‘the processing, recycling, storage, transport and loss of chemical components within the marine environment, mediated by biological processes’. We are said to be leaving the Holocene and entering the ‘Anthropocene’ where mankind’s cumulative impacts have significant and measurable effects on the biosphere. Biogeochemistry lies at the heart of studies on the functioning of marine provinces or types – collectively here referred to as ‘systems’ – that are crucial to understanding and predicting global change and its consequences. In the context of this great environmental and societal impact, it is the varying consequences of the same biogeochemical processes operating in marine systems under different forcing parameters that make biogeochemistry such a diverse and fascinating field. Over the past two decades, much has been learned about the biogeochemical functioning of marine systems from large-scale, multi-partner, international and national research programmes such as are supported by the International Geosphere–Biosphere Program (IGBP), Scientific Committee on Ocean Research (SCOR), Joint Global Ocean Flux Study (JGOFS) and its regional studies, and Land–Ocean Interactions in the Coastal Zone (LOICZ). In the UK, the supporting national programmes were the Biogeochemical Ocean Flux Study (BOFS) and Land–Ocean Interaction Study (LOIS) programmes. These have been undertaken typically on ‘process’ research cruises, where the focus has been on quantifying fluxes of key components (particularly carbon) within the ocean as well as between the ocean and its boundaries (land, sediments and atmosphere). Whilst considerable information continues to be derived from such studies, the expense of such undertakings, together with the relatively low temporal and spatial coverage offered, has led some biogeochemists to develop and use new methods of data collection. These include satellite and airborne remote sensing, benthic landers, autonomous underwater vehicles, and moored and drifting sensor packages with intelligence. Many of these systems have been developed for open ocean deployment, but they are also becoming modified

PREFACE

xiii

for use in shallow, coastal locations. We can expect to see further developments, together with new and more robust sensors and increased data collection and transmission capacity, leading to great improvements in knowledge, operating in a synoptic fashion (for example, the new ARGO programme of drifting subsurface floats across the ocean basins). Modelling has become ubiquitous in biogeochemistry, as in marine science more generally. Significant computing power is now available for the nesting of biogeochemical models within physical oceanographic models with high spatial resolution. Not only does this allow the generalisation of measurements made at a point in space and time, but it allows assessments and comparisons of the relative sensitivities of systems to external changes such as are caused, for example, by increased temperature, deepwater trawling or hydrocarbon exploration. This volume provides an overview of recent research on the biogeochemistry of a diverse range of complex marine systems, each of great importance to the ‘earth system’ but for varying reasons. The systems were chosen to emphasise different forcing factors, thus offering interesting contrasts. We have been fortunate that the chapter authors reflect the diversity of academic backgrounds that typifies biogeochemical research and that they have approached their tasks from varied perspectives. Thus, the repetition of basic concepts between chapters is kept to a minimum. The book will be read by researchers and advanced students of biogeochemistry, who will enjoy the contrasts between the systems chosen, and by workers in related areas of earth science, who will find that it provides a useful point of access to the primary literature across a broad range of marine biogeochemical processes. The first chapter deals with mangroves – key providers of biogeochemical services in large areas of tropical coastal areas that are under threat from insensitive development pressures. We stay in the tropics to consider coral biogeochemistry – also under threat in many areas from a combination of climate change, eutrophication, tourism and destructive fishing – before moving to fjords – the main interface between land and ocean in high latitudes. The eastern Mediterranean continues to attract considerable attention as a highly nutrient-poor and low productivity area, in stark contrast to the Arctic, which is light-limited during the winter months, highly productive in the summer, shows strong benthic–pelagic coupling over the shelf areas and has a productive community associated with the underside of sea ice. In the Arabian Sea, the biogeochemical system is under the control of large-scale, monsoon-linked circulation reversals with a pronounced oxygen minimum zone, and again this is an area under continuing scrutiny for its potential role in the nitrogen biogeochemical cycle. The sediments of the northeast Pacific abyss are dominated by a strong latitudinal gradient of carbon input across the equatorial divergence that has a profound effect on benthic productivity. Even in an area so remote from land, the threat of anthropogenic disturbance in the form of metal nodule mining is very real.

xiv

PREFACE

The penultimate chapter deals with the unique biogeochemistry of the hot and cold vents associated with plate tectonics – largely unknown until relatively recently and now thought to be of great significance in maintaining the basic chemistry of sea water. The book is completed by a modelling section, in which the ecology of planktonic organisms is examined in biogeochemical terms with an emphasis on modelling the interactions between pelagic chemistry and ecology in shelf seas, where significant recycling of sedimentary nutrients supplements direct terrestrial inputs. We are grateful to all the authors who have contributed to this volume. Each author has original insights and approaches and so each chapter is fresh and the whole volume novel and readable. We are particularly indebted to Graeme MacKintosh and David McDade at Blackwell Publishing, who have offered every support and encouragement to this project. Kenneth D. Black Graham B. Shimmield

Contributors

M.J. Atkinson

University of Hawaii SOEST, Hawaii Institute of Marine Biology, PO Box 1346, Kaneohe Bay Hawaii 96744 JoLynn Carroll Akvaplan-niva, Polar Environmental Centre, 14 Hjalmar Johansensgate, N-9296 Tromsø, Norway Michael L. Carroll Akvaplan-niva, Polar Environmental Centre, 14 Hjalmar Johansensgate, N-9296 Tromsø, Norway J.L. Falter University of Hawaii, SOEST, Hawaii Institute of Marine Biology, PO Box 1346, Kaneohe Bay Hawaii 96744 Steven Groom Plymouth Marine Laboratory, Prospect Place, Plymouth PL1 3DH Angelos K. Hannides Department of Oceanography, SOEST, University of Hawaii at Manoa, 1000 Pope Road, Marine Sciences Building, Honolulu, HI 96822 M. Holmer Institute of Biology, University of Southern Denmark, Campusvej 55, DK-5230 Odense, Denmark David Hydes Southampton Oceanography Centre, European Way, Southampton SO14 32H S. Kim Juniper Centre GEOTOP-UQÀM-McGill, Université du Québec à Montréal, C.P. 8888 Succ. Centre-Ville, Montréal, Québec, Canada, H3C 3P8 Michael D. Krom School of Earth Sciences, Leeds University, Leeds LS2 9JT Richard J. Léveillé Centre GEOTOP-UQÀM-McGill, Université du Québec à Montréal, C.P. 8888 Succ. Centre-Ville, Montréal, Québec, Canada, H3C 3P8 B. McKee Tulane University, New Orleans, USA Hema Naik National Institute of Oceanography, Dona Paula, Goa 403 004, India S.W.A. Naqvi National Institute of Oceanography, Dona Paula, Goa 403 004, India P.V. Narvekar National Institute of Oceanography, Dona Paula, Goa 403 004, India

xvi

Richard Sanders J.M. Skei Craig R. Smith

B. Sundby Paul Tett Tamar Zohary

CONTRIBUTORS

Southampton Oceanography Centre, European Way, Southampton SO14 32H Norwegian Institute for Water Research, Oslo, Norway Department of Oceanography, SOEST, University of Hawaii at Manoa, 1000 Pope Road, Marine Sciences Building, Honolulu, HI 96822 ISMER, Université du Québec à Rimouski and McGill University, Montreal, Canada School of Life Sciences, Napier University, 10 Colinton Road, Edinburgh, EH10 5DT Kinneret Limnological Laboratory, Israel Oceanographic and Limnological Research Ltd, PO Box 447, Migdal 14950, Israel

1

Mangroves of Southeast Asia M. Holmer

1.1

Introduction

The last decade has contributed significantly to the development of research on the biogeochemistry of tropical mangrove ecosystems. Also obvious during the last decade is the continued and dramatic destruction of natural tropical mangrove resources in Southeast Asia (Fig. 1.1). The naturally high productivity of tropical mangroves has traditionally been exploited for a wide variety of purposes, both as sources of forestry or fisheries products and they have also been used for human settlement (Hatcher et al., 1989; Platong, 1998). More recently, Southeast Asian mangroves are being extensively cleared for the construction of aquaculture ponds for prawn production (Primavera, 1993). This and other consumptive uses have been estimated to cause an annual reduction of 1% of the world’s tropical mangrove resources (Hatcher et al., 1989). Southeast Asian mangrove forests are declining at alarming rates, due to the increasing demand for land to be allocated to food, industrial production and urban settlements (Kautsky et al., 2000). More than half of the 367 900 ha of mangroves that was present in Thailand in 1961 had already been converted to prawn farms or for various other uses by 1989 (Aksornkoae, 1993), and the mangrove area was further reduced by 81 000 ha in 1996 (Platong, 1998). Mangrove forests in many other Southeast Asian countries also face the same rate of destruction, e.g. Vietnam (Kautsky et al., 2000). Changes in land use in Southeast Asia have resulted in high soil erosion rates and have yielded a major increase in transport of eroded sediments to the coastal zone. The long-term impact and the ramifications of anthropogenic disturbance, such as pollutant discharge, on the biogeochemistry of tropical mangrove forests and their associated near-shore habitats are poorly known. The aim of this chapter is to review recent findings on aspects of the biogeochemistry of Southeast Asian mangrove forests, focusing on physical and biotic processes determining the cycling of elements in mangroves. The chapter will discuss the significance of anthropogenic activities for the biogeochemical cycling of nutrients, which deserves particular attention in the study of Southeast Asian mangrove forests.

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Fig. 1.1 Areas of mangrove (solid line) and mangrove disturbance (hatched area) in the tropics.

1.2

Mangrove forest structure and function

Earlier models of tropical mangrove forests proposed overwhelming influences by physical forces and processes such as the tidal regime or geomorphology on ecosystem structure and function. Mangrove ecosystem development was depicted as successional systems, where the presence of plants themselves had a significant impact on the physical environment; and such impact culminated in an alternation of growth conditions favoured by different species in time. In addition, recent findings (especially from the Indo-Pacific mangrove forests) suggest considerable influence of biotic agents and processes such as sesarmine crab feeding and bioturbation activities in shaping the ecology of tropical mangrove forests (Twilley et al., 1997). Biotic influences on mangrove forest ecosystem structure and function are expected to be more important in systems with weak external forcing or high biodiversity. Climatic features, such as the timing of monsoon periods, can give rise to strong seasonality (Alongi & Sasekumar, 1992). Most productive mangrove ecosystems are highly effective sinks for nutrients essential to sustain high rates of plant growth, as evidenced from the fact that many such systems export refractory, particulate organic carbon but import most dissolved nutrient species (Alongi et al., 1992; Robertson et al., 1992; Hemminga et al., 1994; Rivera-Monroy et al., 1995; Alongi, 1996). Several studies have suggested that close couplings exist among benthic nutrient pools,

MANGROVES OF SOUTHEAST ASIA

3

microbes and mangrove trees, acting as mechanisms to maximize utilization and conservation of scarce nutrients (Boto & Wellington, 1983, 1984; Kristensen et al., 1995; Alongi, 1996).

1.3

Water column biogeochemistry

Mangrove forests are highly diversified due to the large variability, for example, in geomorphology and tidal activities, and a number of different functional types of mangroves have been described (Woodroffe, 1992). The hydrodynamics in some mangrove forests are strongly influenced by river inputs, whereas others are much more dominated by the ocean, as found for fringing mangrove forests. A large number of local factors may thus influence the water column processes, which makes it very difficult to provide a generalized description of the water biogeochemistry in mangrove forests. Mangrove creeks are considered as important routes for tidal exchange of dissolved and particulate matter between the forest environment and adjacent coastal waters (Wolanski et al., 1992; Hemminga et al., 1994; Rivera-Monroy et al., 1995). The residence time of the water in the creeks is usually a useful indicator of the biogeochemical fate of the compounds in the water column (Suraswadi et al., 2003), where long residence times allow for uptake of nutrients by the vegetation, e.g. phytoplankton and mangrove trees, while short residence times leads to a larger export of materials to the ocean (Thong et al., 1993). Hydrodynamics in the mangrove forest is controlled by tides, mangrove vegetation and geometry of the mangrove waterways (Hoguane et al., 1999). Friction from dense mangrove trees influences the tidal regime and causes tidal asymmetry (Suraswadi et al., 2003), and current velocities in a channel cross section have lateral or vertical variations due to channel geometry and bathymetry (Valle-Levinson & Atkinson, 1999). These variations in water velocity cause transverse and vertical shear stress, which are important for the mixing process in the creek water (Uncle et al., 1985). The residence time of water in mangrove forests is quite variable, determined by the forest topography, size and type, and thus hydrodynamics. It can vary from a few days in small fringe forests exposed to large tidal variation (Wattayakorn et al., 1990) to more than a month in large mangrove forests (Wolanski et al., 1990). Suraswadi et al. (2003) studied the hydrodynamics in a small mangrove forest in Thailand. The hydrodynamics was modified as a result of friction created by the mangrove vegetation, and these modifications resulted in strong ebb current, asymmetric flood and ebb tide and a time lag in the tidal phase between the upper and lower creek. The main creek was well mixed with a transient stratification during low tide and was completely mixed during high tide. This situation is similar to other mangrove estuaries (Wattayakorn et al., 1990; Wolanski et al., 1990). The hydrodynamics may be influenced by heavy rains; in this mangrove, rainfall caused an increased transport of salt water when the

4

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rainwater pushed the creek water outward, and the period and amount of rainfall must thus be considered when the flux of solutes from the mangrove forest is determined during the wet season. Nutrient levels in pristine tropical mangrove forests vary both in time and space as a result of differences in hydrodynamics, freshwater input, solar insolation, and productivity of phytoplankton and bacterioplankton (Ovalle et al., 1990; Alongi et al., 1992; Bava & Seralathan, 1998; Trott & Alongi, 1999; Ayakai et al., 2000). Mangrove creeks are, however, usually characterized by low nutrient concentrations due to a high capacity for retaining and recycling of nutrients within the system (Kristensen et al., 1995). Even in areas with high nutrient loading, e.g. due to urban settlement, nutrient levels are generally low (Harrison et al., 1997). Nutrient cycling in the water column is controlled by a large number of auto- and heterotrophic processes (Fig. 1.2), and the dissolved

Fig. 1.2 Nutrient transformation processes in the water column in mangrove forests.

MANGROVES OF SOUTHEAST ASIA

5

organic matter pool especially is considered to play an important role (Alongi et al., 1989; Bano et al., 1997; Ayukai et al., 1998). Leakage of nutrient rich water from the creek banks during low tides has been suggested as an important contributor to nutrients in the mangrove waterways, and was investigated in a mangrove forest in Thailand (Kristensen & Suraswadi, 2002). Water seeping from creek banks was only enriched in inorganic phosphates and was not considered as an important source of solutes to the waterways in this forest. The low nutrient concentrations affect the primary production in mangrove creeks, and the production has often been found to be strongly nutrient limited. Phosphorus was the limiting factor for phytoplankton growth in a mangrove creek in a tidal-dominated forest in Thailand, as indicated by a high molar ratio between dissolved nitrogen and phosphorus (~34–38) much higher than the Redfield ratio (Suraswadi et al., 2003). The degree of phosphorus limitation was less pronounced near small tributaries (~11–21), whereas the N:P ratio increased significantly (~192) near shrimp farm outlets – most likely in response to loss of dissolved nitrogen compounds from the farms (Burford & Longmore, 2001). The phytoplankton production is, however, not always nutrient limited. Due to the shallow water in many mangrove creeks and a rapid water flow in river-influenced mangrove forests the light penetration is often quite low and the phytoplankton production is just as often light limited (Harrison et al., 1997; Kristensen & Suraswadi, 2002). In a three year study of two tidal creeks on the Indus River in Pakistan, Harrison et al. (1997) found no limitation of the phytoplankton production by silicate or nitrogen, and phosphate was only limiting during large blooms. Conversely a large suspended load resulted in high light extinction coefficients and the 1% light depth ranged from > kSi,j is: μ .τ u maxi, j 1 Q i, j, t = 0 ⎞ ⎛⎛e t i . B maxi ≈ B i, t = 0 ⋅ ⎜ ---------- ⋅ k i, j1 ⋅ --------------⎞ + -----------------⎝ μi ⎠ k k Qi ⋅ j ⎟⎠ Qi, j1 ⎝

(7)

The parameter τi, gives the time at which the ambient nutrient concentration falls to that of the uptake half-saturation concentration for (limiting) nutrient j1. Population growth during this period is assumed to take place at a constant rate given by μi (d−1), more likely limited by light than by nutrient. The term ki takes account of the mean value of S/ks+S during this period, which is (by definition of the conditions) between 0.5 and 1. Mainly, however, it is influenced by the value of finhib (Qi). In effect, a species, or life-form, that grows with Q substantially less than Qmax, has a high value of k, which thus

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give its capability for luxury uptake. Thus, success amongst species in this yield-limited case is determined by the matrix of values of ki,j, umaxi,j and μi as well as those of kQi,j. The second class of solutions involves a steady state, which might occur in a laboratory chemostat, or in the upper waters of a tropical ocean where losses Li = μi are due to physical removal and grazing. In the steady state, ui,j = ρi,j ·μi · Qi,j. Given Sj 1 implies nutrient 1 limiting, and ρ* < 1 implies nutrient 2 limiting. S j 1 k Sj 1 u maxj 1 k Qj 2 Nutrient 1 is limiting when: ------ < ------- ⋅ ------------- ⋅ -------- . A solution for a limitS j2 k Sj2 u maxj2 k Qj1 ing nutrient and several species is: ℜ=

u maxi

- ⋅ B i⎞ ∑ ( u i ⋅ B i ) ≈ S ⋅ ∑ ⎛⎝ ---------⎠ kS i

i

(9)

i

where ℜ is the input of the nutrient (g-at m−3 d−1) due to vertical mixing and local remineralisation. In this uptake-limited case, the available nutrient supply is distributed amongst biota in proportion to their biomasses and their ratio of maximum uptake rate to half-saturation concentration. However, the steady state is unstable, as a small increase in biomass results in that species getting more of the available nutrient, leaving less for others: in principle the outcome is that all Bi except one become zero. Other species can survive if they are limited by different nutrients. Species limited by the same nutrient may also co-exist if grazing keeps their biomasses near a grazing threshold Bthr such that each population’s nutrient demand μi · Qi,j ·Bthr is much less than ℜ. Dedication Paul Tett wishes to dedicate this chapter to the memory of Mahlon G. Kelly, late of the University of Virginia, who introduced him to Biogeochemistry.

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References Aksnes, D.L., Egge, J.K., Rosland, R. and Heimdal, B.R. (1994) Representation of Emiliania huxleyi in phytoplankton simulation-models – a 1st approach. Sarsia, 79, 291–300. Allen, J.R., Slinn, D.J., Shammon, T.M., Hartnoll, R.G. and Hawkins, S.J. (1998) Evidence for eutrophication of the Irish Sea over four decades. Limnology and Oceanography, 43, 1970–1974. Asman, W.A.H., Hertel, O., Bercowicz, R., Christensen, J., Runge, E.H., Sorensen, L.L., Granby, K., Nielsen, H., Jensen, B., Gryning, S.E., Sempreviva, A.M., Larsen, S., Hummelshoj, P., Jensen, N.O., Allerup, P., Jorgensen, J., Madsen, H., Overgaard, S. and Vejen, F. (1995) Atmospheric nitrogen input to the Kattegat. Ophelia, 42, 5–28. Azam, F., Fenchel, T., Field, J.G., Gray, J.S., Meyer-Reil, L.A. and Thingstad, F. (1983) The ecological role of water-column microbes in the sea. Marine Ecology Progress Series, 10, 257–263. de Baar, H.J.W. and de Jong, J.T.M. (2001) Distributions, sources and sinks of iron in seawater, in The Biogeochemistry of iron in seawater (eds D.R. Turner and K.A. Hunter), pp. 123–253. Bates, S.S., de Freitas, A.S.W., Milley, J.E., Pocklington, R., Quilliam, M.A., Smith, J.C. and Worms, J. (1991) Controls on domoic acid production by the diatom Nitzschia pungens f. multiseries in culture: nutrients and irradiance. Canadian Journal of Fisheries and Aquatic Science, 48, 1136–1144. Billen, G., Lancelot, C. and Meybeck, M. (1991) N, P and Si retention along the aquatic continuum from land to ocean, in Ocean Margin Processes in Global Change (eds R.F.C. Mantoura, J.-M. Martin and R. Wollast), John Wiley and Sons Ltd, pp. 19–44. Billen, G., Somville, M., De Becker, E. and Servais, P. (1985) A nitrogen budget of the Scheldt hydrographic basin. Netherlands Journal of Sea Research, 19, 223–230. Bowman, M.J., Esias, W.E. and Schnitzer, M.B. (1981) Tidal stirring and the distribution of phytoplankton in Long Island and Block Island Sounds. Journal of Marine Research, 39, 587–603. Brand, L.E. (1991) Minimum Iron requirements of marine phytoplankton and the implications for the biogeochemical control of new production. Limnology and Oceanography, 36, 1756–1771. Brockmann, U.H. and Kattner, G. (1997) Winter-to-summer changes of nutrients, dissolved and particulate organic material in the North Sea. Deutsche Hydrographische Zeitschrift, 49, 229–242. Brockmann, U.H. and Wegner, G. (1995) Distribution of nutrients in the central North Sea. Deutsche Hydrographische Zeitschrift, Supplement, 5, 237–249. Bruland, K.W., Donat, J.R. and Hutchins, D.A. (1991) Interactive influences of bioactive trace metals on biological production in oceanic waters. Limnology and Oceanography, 36, 1555–1577. Bruland, K.W., Rue, E.L. and Smith, G.J. (2001) Iron and macronutrients in California coastal upwelling regimes: Implications for diatom blooms. Limnology and Oceanography, 46, 1661–1674. Brzezinski, M.A. (1985) The Si:C:N ratio of marine diatoms: interspecific variability and the effect of some environmental variables. Journal of Phycology, 21, 347–357. Caron, D.A., Goldman, J.C. and Dennett, M.R. (1990) Carbon utilisation by the omnivorous flagellate Paraphysomonas imperforata. Limnology and Oceanography, 35, 192–201. Caron, D.A., Lim, E.L., Sanders, R.W., Dennett, M.R. and Berninger, U.-G. (2000) Responses of bacterioplankton and phytoplankton to organic carbon and inorganic nutrient additions in contrasting oceanic ecosystems. Aquatic Microbial Ecology, 22, 175–184. Carr, N.G. and Wyman, M. (1986) Cyanobacteria: their biology in relation to oceanic picoplankton. Photosynthetic picoplankton, in Canadian Bulletin of Fisheries and Aquatic Sciences, vol. 214 (eds T. Platt and W.K.W. Li), pp. 159–204. C.E.C. (1991) Council Directive of 21 May 1991 concerning urban waste water treatment (91/271/ EEC). Official Journal of the European Communities, L135 of 30.5.91, 40–52. Conley, D.J., Schelske, C.L. and Stoermer, E.F. (1993) Modification of the biogeochemical cycle of silica with eutrophication. Marine Ecology Progress Series, 101, 179–192. Copin-Montegut, C. and Copin-Montegut, G. (1983) Stochiometry of carbon, nitrogen and phosphorus in marine particulate matter. Deep-Sea Research, 30, 31–46.

352

BIOGEOCHEMISTRY OF MARINE SYSTEMS

Croot, P.L., Karlson, B., Wulff, A., Linares, F. and Andersson, K. (2002) Trace metal/phytoplankton interactions in the Skagerrak. Journal of Marine Systems, 35, 39–60. Currie, D.J. and Kalff, J. (1984) A comparison of the abilities of freshwater algae and bacteria to aquire and retain phosphorus. Limnology and Oceanography, 29, 298–310. Davidson, K., Cunningham, A. and Flynn, K.J. (1995) Predator–prey interactions between Isochrysis galbana and Oxyrrhis marine. III. Mathematical modelling of predation and nutrient regeneration. Journal of Plankton Research, 17, 465–492. Dippner, J.W. (1998) Competition between different groups of phytoplankton for nutrients in the southern North Sea. Journal of Marine Systems, 14, 181–198. Droop, M.R. (1968) Vitamin B12 and marine ecology. IV. The kinetics of uptake, growth and inhibition in Monochrysis lutheri. Journal of the Marine Biological Association of the United Kingdom, 48, 689–733. Droop, M.R. (1973) Some thoughts on nutrient limitation in algae. Journal of Phycology, 9, 264–272. Droop, M.R. (1979) On the definition of X and of Q in the Cell Quota model. Journal of Experimental Marine Biology and Ecology, 39, 203. Droop, M.R. (1983) 25 years of algal growth kinetics – a personal view. Botanica Marina, 26, 99–112. Ducobu, H., Huisman, J., Jonker, R.R. and Mur, L.R. (1998) Competition between a prochlorophyte and a cyanobacterium under various phosphorus regimes: Comparison with the Droop model. Journal of Phycology, 34, 467–476. Dugdale, R.C. (1967) Nutrient limitation in the sea: dynamics, identification, and significance. Limnology and Oceanography, 12, 685–695. Dugdale, R.C. and Goering, J.J. (1967) Uptake of new and regenerated forms of nitrogen in primary productivity. Limnology and Oceanography, 12, 196–206. Dussart, B.M. (1965) Les différentes catégories de plancton. Hydrobiologia, 26, 72–74. Dyer, K.R. and Moffat, T.J. (1998) Fluxes of suspended matter in the East Anglian plume, Southern North Sea. Continental Shelf Research, 18, 1311–1331. Egge, J.K. (1998) Are diatoms poor competitors at low phosphate concentrations? Journal of Marine Systems, 16, 191–198. Egge, J.K. and Aksnes, D.L. (1992) Silicate as regulating nutrient in phytoplankton competition. Marine Ecology Progress Series, 83, 281–289. Egge, J.K. and Heimdal, B.R. (1994) Blooms of phytoplankton including Emiliania huxleyi (Haptophyta) – effects of nutrient supply in different N-P ratios. Sarsia, 79, 333–348. Egge, J.K. and Jacobsen, A. (1997) Influence of silicate on particulate carbon production in phytoplankton. Marine Ecology Progress Series, 147, 219–230. Eisma, D. and Kalf, J. (1987) Dispersal, concentration and deposition of suspended matter in the North Sea. Journal of the Geological Society, 144, 161–178. Elliott, J.A., Tett, P., Reynolds, C.S. and Irish, A.E. (2002) Adapting a freshwater model (PROTECH) to simulate phytoplankton in a Scottish sea-loch. Journal of Marine Research, submitted June 2002. Elmgren, R. and Larsson, U. (eds) (1997) Himmerfjärden, Stockholm, Naturvårdsverket Förlag. Elmgren, R. and Larrson, U. (2001) Nitrogen and the Baltic Sea: managing nitrogen in relation to phosphorus. Optimising Nitrogen Management in Food and Energy Production and Environmental protection: Proceedings of the 2nd International Nitrogen Conference on Science and Policy. The Scientific World, 1(S2), 371–377. Engqvist, A. (1996) Long-term nutrient balances in the eutrophication of the Himmerfjarden estuary. Estuarine, Coastal and Shelf Science, 42, 483–507. Eppley, R.W. and Peterson, B.J. (1979) Particulate organic matter flux and planktonic new production in the deep ocean. Nature, 282, 677–680. Escaravage, V., Prins, T.C., Nijdam, C., Smaal, A.C. and Peeters, J.C.H. (1999) Response of phytoplankton communities to nitrogen input reduction in mesocosm experiments. Marine Ecology Progress Series, 179, 187–199.

INFLUENCE OF NUTRIENT BIOGEOCHEMISTRY ON ECOLOGY

353

Fenchel, T. (1984) Suspended marine bacteria as food source, in Flows of energy and materials in marine ecosystems (ed. M.J.R. Fasham), Plenum Press, pp. 301–315. Fichez, R., Dennis, P., Fontaine, M.F. and Jickells, T.D. (1993) Isotopic and biochemical composition of particulate organic matter in a shallow water estuary (Great Ouse, North Sea, England). Marine Chemistry, 43, 263–276. Fichez, R., Jickells, T.D. and Edmunds, H.M. (1992) Algal blooms in high turbidity, a result of the conflicting consequences of turbulence on nutrient cycling in a shallow water estuary. Estuarine, Coastal and Shelf Science, 35, 577–592. Finni, T., Kononen, K., Olsonen, R. and Wallstrom, K. (2001) The history of cyanobacterial blooms in the Baltic Sea. Ambio, 30, 172–178. Flynn, K. (2001) A mechanistic model for describing dynamic multi-nutrient, light, temperature interactions in phytoplankton. Journal of Plankton Research, 23, 977–997. Flynn, K.J. and Hipkin, C.R. (1999) Interactions between iron, light, ammonium and nitrate: insights from the construction of a dynamic model of algal physiology. Journal of Phycology, 35, 1171–1190. Flynn, K.J., Page, S., Wood, G. and Hipkin, C.R. (1999) Variations in the maximum transport rates for ammonium and nitrate in the prymnesiophyte Emiliania huxleyi and the raphidophyte Heterosigma carterae. Journal of Plankton Research, 21, 355–371. Gambin, F., Boge, G. and Jamet, D. (1999) Alkaline phosphatase in a littoral Mediterranean marine ecosystem: role of the main plankton size. Marine Environmental Research, 47, 441–456. Geider, R.J. and La Roche, J. (2002) Redfield revisited: variability of C:N:P in marine microalgae and its biochemical basis. European Journal of Phycology, 37, 1–17. Gieskes, W.W.C. and Kraay, G.W. (1984) Phytoplankton, its pigments, and primary production at a central North Sea station in May, July and September 1981. Netherlands Journal of Sea Research, 18, 51–70. Gillbricht, M. (1988) Phytoplankton and nutrients in the Helgoland region. Helgolander Meeresuntersuchungen, 42, 435–467. Gilpin, L.C., Davidson, K. and Roberts, E.C. The influence of changes in nitrogen:silicon ratios on diatom growth dynamics. Journal of Sea Research (accepted, 2003). Gismervik, I., Andersen, T. and Vadstein, O. (1996) Pelagic food webs and eutrophication of coastal waters: impact of grazers on algal communities. Marine Pollution Bulletin, 33, 22–35. Gledhill, M. and van den Berg, C.M.G. (1994) Determination of complexation of iron(III) with natural organic complexing ligands in sea water using cathodic stripping voltametry. Marine Chemistry, 47, 41–54. Gledhill, M., van den Berg, C.M.G., Nolting, R.F. and Timmermans, K.R. (1998) Variability in the speciation of iron in the northern North Sea. Marine Chemistry, 59, 283–300. Goldman, J.C. and Dennett, M.R. (1991) Ammonium regeneration and carbon utilization by marine bacteria grown on mixed substrates. Marine Biology, 109, 369–378. Goldman, J.C., Caron, D.A. and Dennett, M.R. (1987) Regulation of gross growth efficiency and ammonium regeneration in bacteria by substrate C:N ratio. Limnology and Oceanography, 32, 1239–1252. Gowen, R.J., Hydes, D.J., Mills, D.K., Stewart, B.M., Brown, J., Gibson, C.E., Shammon, T.M., Allen, M. and Malcolm, S.J. (2002) Assessing trends in nutrient concentrations in coastal shelf seas: a case study in the Irish Sea. Estuarine, Coastal and Shelf Science, 54, 927–939. Gowen, R.J., Mills, D.K., Trimmer, M. and Nedwell, D.B. (2000) Production and its fate in two coastal regions of the Irish Sea: the influence of anthropogenic nutrients. Marine Ecology Progress Series, 208, 51–64. Gran, H.H. (1931) On the conditions for the production of plankton in the sea. Rapports et ProcesVerbaux des Reunions du Conseil International pour l’Exploration de la Mer, 75, 37–46. Granéli, E. and Risinger (1994) Effects of Cobalt and Vitamin B12 on the growth of Chrysochromulina polylepis (Prymnesiophyceae). Marine Ecology Progress Series, 113, 177–183.

354

BIOGEOCHEMISTRY OF MARINE SYSTEMS

Greene, R.M., Geider, R.J. and Falkowski, P.G. (1991) Effect of iron limitation on photosynthesis in a marine diatom. Limnology and Oceanography, 36, 1772–1782. Grice, G.D., Harris, R.P., Reeve, M.R., Heinbokel, J.F. and Davis, C.O. (1980) Large-scale enclosed water column ecosystems: an overview of Foodweb I, the final CEPEX experiment. Journal of the Marine Biological Association of the United Kingdom, 60, 401–414. Grimes, J.P. (1979) Plant strategies and vegatative processes, Wiley, Chichester. Hamm, C.E., Simson, D.A., Merkel, R. and Smetacek, V. (1999) Colonies of Phaeocystis globosa are protected by a thin but tough skin. Marine Ecology Progress Series, 187, 101–111. Hansen, B., Bjørnsen, P.K. and Hansen, P.J. (1994) The size ratio between planktonic predators and their prey. Limnology and Oceanography, 39, 395–403. Hansen, P.J. (1991) Quantitative importance and trophic role of heterotrophic dinoflagellates in a coastal pelagial food web. Marine Ecology Progress Series, 73, 253–261. Hansen, P.J., Nielsen, T.G. and Kaas, H. (1995) Distribution and growth of protists and mesozooplankton during a bloom of Chrysochromulina spp (Prymnesiophyceae, Prymnesiales). Phycologia, 34, 409–416. Harada, S., Watanabe, M., Kohata, K., Ioriya, T., Kunugi, M., Kimura, T., Fujimori, S., Koshikawa, H. and Sato, K. (1996) Analyses of planktonic ecosystem structure in coastal seas using a largescale stratified mesocosm: a new approach to understanding the effects of physical, biochemical and ecological factors on phytoplankton species succession. Water Science and Technology, 34, 219–226. Hardy, A.C. (1924) The herring in relation to its animate environment. Part I. The food and feeding habits of the herring with special reference to the east coast of England. Fishery Investigations, Series II, 7(3), 1–53. Harrison, G.I. and Morel, F.M.M. (1986) Response of the marine diatom Thalassiosira weisflogii to iron stress. Limnology and Oceanography, 31, 989–997. Harrison, P.J., Conway, H.L. and Dugdale, R.C. (1976) Marine diatoms grown in chemostats under silicate or ammonium limitation. I. Cellular chemical composition and steady-state growth kinetics of Skeletonema costatum. Marine Biology, 35, 177–186. Hastings, A. (1996) Population Biology: concepts and models, Springer, New York. Head, R.N., Crawford, D.W., Egge, J.K., Harris, R.P., Kristiansen, S., Lesley, D.J., Maranon, E., Pond, D. and Purdie, D.A. (1998) The hydrography and biology of a bloom of the coccolithophorid Emiliania huxleyi in the northern North Sea. Journal of Sea Research, 39, 255–266. Hecky, R.E. and Kilham, P. (1988) Nutrient limitation of phytoplankton in freshwater and marine environments: a review of recent evidence on the effects of enrichment. Limnology and Oceanography, 33, 796–822. Henriksen, P., Knipschildt, F., Moestrup, O. and Thomsen, H.A. (1993) Autecology, life-history and toxicology of the silicoflagellate Dictyocha speculum (Silicoflagellata, Dictyochophyceae). Phycologia, 32, 29–39. Hickel, W. (1998) Temporal variability of micro- and nanoplankton in the German Bight in relation to hydrographic structure and nutrient changes. ICES Journal of Marine Science, 55, 600–609. Hickel, W., Mangelsdorf, P. and Berg, J. (1993) The human impact in the German Bight – eutrophication during 3 decades (1962–1991). Helgolander Meeresuntersuchungen, 47, 243–263. Hill, A.E. and Simpson, J.H. (1988) Low-frequency variability of the Scottish Coastal Current induced by along-shore pressure gradients. Estuarine, Coastal and Shelf Science, 27, 163–180. Hobson, L.A. (1989) Paradox of the phytoplankton – an overview. Biological Oceanography, 6, 493–504. Hodgkiss, I.J. and Ho, K.C. (1997) Are changes in N:P ratios in coastal waters the key to increased red tide blooms? Hydrobiologia, 352, 141–147. Holligan, P.M. (1981) Biological implications of fronts on the northwest European continental shelf. Philosophical Transactions of the Royal Society of London Series a Mathematical, Physical and Engineering Sciences, 302, 547–562. Holligan, P.M., Harris, R.P., Newell, R.C., Harbour, D.S., Head, R.N., Linley, E.A., Lucas, M.I., Tranter, P.R.G. and Weekly, C.M. (1984) Vertical distribution and partitioning of organic carbon

INFLUENCE OF NUTRIENT BIOGEOCHEMISTRY ON ECOLOGY

355

in mixed, frontal and stratified waters of the English Channel. Marine Ecology Progress Series, 14, 111–127. Howarth, M.J., Dyer, K.R., Joint, I.R., Hydes, D.J., Purdie, D.A., Edmunds, H., Jones, J.E., Lowry, R.K., Moffat, T.J., Pomroy, A.J. and Proctor, R. (1993) Seasonal cycles and their spatial variability. Philosophical Transactions of the Royal Society of London Series a Mathematical, Physical and Engineering Sciences, 343, 383–403. Hutchins, D.A., Franck, V.M., Brzezinski, M.A. and Bruland, K.W. (1999) Inducing phytoplankton iron limitation in iron-replete coastal waters with a strong chelating ligand. Limnology and Oceanography, 44, 1009–1018. Hutchins, D.A., Sedwick, P.N., DiTullio, G.R., Boyd, P.W., Queguiner, B., Griffiths, F.B. and Crossley, C. (2002) Control of phytoplankton growth by iron and silicic acid availability in the subantarctic Southern Ocean: Experimental results from the SAZ Project. Journal Of Geophysical Research – Oceans, 106, 31559–31572. Hutchinson, G.E. (1961) The paradox of the plankton. American Naturalist, 93, 137–145. Huthnance, J.M. (1995) Circulation, exchange and water masses at the ocean margin: the role of physical processes at the shelf edge. Progress in Oceanography, 35, 353–431. Hydes, D.J. and Edmunds, H. (1992) Nutrients in the North Sea, September 1988 to October 1989: use of nutrient ratios and nutrient salinity relationships to identify sources and sinks of nutrients. Science of the Total Environment, Supplement 1992, 1149–1157. Hydes, D.J., Kelly-Gerreyn, B.A., Le Gall, A.C. and Proctor, R. (1999) The balance of supply of nutrients and the demands of biological production and denitrification in a temperate latitude shelf sea – a treatment of the southern North Sea as an extended estuary. Marine Chemistry, 68, 117–131. Hydes, D.J., Le Gall, A.C., Miller, A.E.J., Brockmann, U., Rabbe, T., Holley, S., Alvarez-Salgado, X., Antia, A., Balzer, W., Chou, L., Elskens, M., Helder, W., Joint, I. and Orren, M. (2001) Supply and demand of nutrients and dissolved organic matter at and across the N.W. European shelf break in relation to hydrography and biogeochemical activity. Deep-Sea Research II – Topical Studies In Oceanography, 48, 3023–3047. Irigoien, X., Harris, R.P., Head, R.N. and Harbour, D. (2000) The influence of diatom abundance on the egg production rate of Calanus helgolandicus in the English Channel. Limnology and Oceanography, 45, 1433–1439. Istvanovics, V., Shafik, H.M., Presing, M. and Juhos, S. (2000) Growth and phosphate uptake kinetics of the cyanobacterium, Cylindrospermopsis raciborskii (Cyanophyceae) in throughflow cultures. Freshwater Biology, 43, 257–275. Jacobsen, A., Egge, J.K. and Heimdal, B.R. (1995) Effects of increased concentration of nitrate and phosphate during a spring bloom experiment in mesocosm. Journal of Experimental Marine Biology and Ecology, 187, 239–251. Jeffrey, S.W. and Vesk, M. (1997) Introduction to marine phytoplankton and their pigment signatures, in Phytoplankton pigments in oceanography: guidelines to modern methods (eds S.W. Jeffrey, R.F.C. Mantoura and S.W. Wright), UNESCO, Paris, Monographs on Oceanographic Methodology. Jickells, T. (1995) Atmospheric inputs of metals and nutrients to the oceans: Their magnitude and effects. Marine Chemistry, 48, 199–214. Jickells, T., Andrews, J., Samways, G., Sanders, R. and Malcolm, S. (2000) Nutrient fluxes through the Humber estuary – past, present and future. Ambio, 29, 130–135. Jickells, T.D. (1998) Nutrient biogeochemistry of the coastal zone. Science, 281, 217–222. Johansson, N. and Granéli, E. (1999) Cell density, chemical composition and toxicity of Chrysochromulina polylepis (Haptophyta) in relation to different N:P supply ratios. Marine Biology, 135, 209–217. Johnson, K.S., Gordon, R.M. and Coale, K.H. (1997) What controls dissolved iron concentrations in the world ocean? Marine Chemistry, 57, 137–161. Joint, I. and Pomroy, A. (1993) Phytoplankton biomass and production in the southern North Sea. Marine Ecology Progress Series, 99, 169–182.

356

BIOGEOCHEMISTRY OF MARINE SYSTEMS

Joint, I.R. (1986) Physiological Ecology of picoplankton in various oceanographic provinces, in Photosynthetic picoplankton. Canadian Bulletin of Fisheries and Aquatic Sciences, vol. 214 (eds T. Platt and W.K.W. Li), pp. 287–309. Jones, H.L.J., Leadbeater, B.S.C. and Green, J.C. (1993) Mixotrophy in marine species of Chrysochromulina (Prymnesiophyceae) – ingestion and digestion of a small green flagellate. Journal of the Marine Biological Association of the United Kingdom, 73, 283–296. Jones, K.J. (1979) Studies on nutrient levels and phytoplankton growth in a Scottish Sea Loch. PhD thesis, University of Strathclyde, pp. 390. Jones, K.J. and Gowen, R.J. (1990) Influence of stratification and irradiance regime on summer phytoplankton composition in coastal and shelf seas of the British Isles. Estuarine, Coastal and Shelf Science, 30, 557–567. Jones, K.J., Gowen, R.J. and Tett, P. (1984) Water-column structure and summer phytoplankton distribution in the Sound of Jura. Journal of Experimental Marine Biology and Ecology, 78, 269–289. Justic, D., Rabalais, N.N. and Turner, R.E. (1995) Stoichiometric nutrient balance and origin of coastal eutrophication. Marine Pollution Bulletin, 30, 41–46. Katechakis, A., Stibor, H., Sommer, U. and Hansen, T. (2002) Changes in the phytoplankton community and microbial food web of Blanes Bay (Catalan Sea, NW Mediterranean) under prolonged grazing pressure by doliolids (Tunicata), cladocerans or copepods (Crustacea). Marine Ecology Progress Series, 234, 55–69. Kawachi, M., Inouye, I., Maeda, O. and Chihara, M. (1991) The haptonema as a food-capturing device – observations on Chrysochromulina hirta (Prymnesiophyceae). Phycologia, 30, 563–573. Kelly, M.G. and McGrath, J.C. (1975) Biology: Evolution and Adaptation to the Environment, Houghton Mifflin, Boston, Mass. Kiørboe, T. (1993) Turbulence, phytoplankton cell size and the structure of pelagic food webs. Advances in Marine Biology, 29, 1–72. Knight, P.J. and Howarth, M.J. (1999) The flow through the north channel of the Irish Sea. Continental Shelf Research, 19, 693–716. Koehl, M.A.R. (1984) Mechanisms of particle capture by copepods at low Reynolds number: possible modes of selective feeding, in Trophic interactions within aquatic ecosystems (eds D.G. Meyers and J.R. Strickler), American Association for the Advancement of Science, Washington, pp. 135–166. Kuma, K., Nishioka, J. and Matsunaga, K. (1996) Controls on iron(III) hydroxide solubility in sea water: influence of pH and natural organic chelators. Limnology and Oceanography, 41, 396–407. Laane, R.W.P.M., Groeneveld, G., Devries, A., van Bennekom, J. and Sydow, S. (1993) Nutrients (P, N, Si) in the Channel and the Dover Strait – seasonal and year-to-year variation and fluxes to the North Sea. Oceanologica Acta, 16, 607–616. Lancelot, C. (1990) Phaeocystis blooms in the continental coastal area of the Channel and the North Sea, in Eutrophication and algal blooms in North Sea coastal zones, the Baltic and adjacent areas: prediction and assessment of preventive actions (eds C. Lancelot, G. Billen and H. Barth) Commission of the European Communities, Brussels, Water Pollution Research Report, 12, pp. 27–54. Lancelot, C. and Billen, G. (1985) Carbon–nitrogen relationships in nutrient metabolism of coastal marine ecosystems. Advances in Aquatic Microbiology, 3, 263–321. Lancelot, C., Billen, G., Sournia, A., Weisse, T., Colijn, F., Veldhuis, M.J.W., Davies, A. and Wassman, P. (1987) Phaeocystis blooms and nutrient enrichment in the continental coastal zones of the North Sea. Ambio, 16, 38–46. Larrson, U., Elmgren, R. and Wulff, F. (1985) Eutrophication and the Baltic Sea – causes and consequences. Ambio, 14, 9–14. Lauria, M.L., Purdie, D.A. and Sharples, J. (1999) Contrasting phytoplankton distributions controlled by tidal turbulence in an estuary. Journal of Marine Systems, 21, 189–197. Lazier, J.R.N. and Mann, K.H. (1989) Turbulence and the diffusive layer around small organisms. Deep-Sea Research, 36, 1721–1733.

INFLUENCE OF NUTRIENT BIOGEOCHEMISTRY ON ECOLOGY

357

Lee, J.-Y., Tett, P., Jones, K., Jones, S., Luyten, P., Smith, C. and Wild-Allen, K. (2002) The PROWQM physical-biological model with benthic-pelagic coupling applied to the northern North Sea. Journal of Sea Research, 48, 287–331. Le Gall, A.C., Hydes, D.J., Kelly-Gerreyn, B. and Slinn, D.J. (2000) Development of a 2D horizontal biogeochemical model for the Irish Sea DYMONIS. ICES Journal of Marine Science, 57, 1050–1059. Legendre, L. and Rassoulzadegan, F. (1995) Plankton and nutrient dynamics in marine waters. Ophelia, 41, 153–172. Lewis, J. (1988) Cysts and sediments: Gonyaulax polyedra (Lingulodinium machaerophorum) in Loch Creran. Journal of the Marine Biological Association of the United Kingdom, 68, 701–714. Liu, H.B., Laws, E.A., Villareal, T.A. and Buskey, E.J. (2001) Nutrient-limited growth of Aureoumbra lagunensis (Pelagophyceae), with implications for its capability to outgrow other phytoplankton species in phosphate-limited environments. Journal of Phycology, 37, 500–508. Lohse, L., Kloosterhuis, H.T., van Raaphorst, W. and Helder, W. (1996) Denitrification rates as measured by the isotope pairing method and by the acetylene inhibition technique in continental shelf sediments of the North Sea. Marine Ecology Progress Series, 132, 169–179. Maestrini, S.Y. and Granéli, E. (1991) Environmental conditions and ecophysiological mechanisms which led to the 1988 Chrysochromulina polylepis bloom – an hypothesis. Oceanologica Acta, 14, 397–413. Maestrini, S.Y., Béchmein, C., Grzebyk, D. and Hummert, C. (2000) Phosphorus limitation might promote more toxin content in the marine invader dinoflagellate Alexandrium minutum. Plankton Biology and Ecology, 47, 7–11. Maldonado, M.T. and Price, N.M. (1999) Utilization of iron bound to strong organic ligands by plankton communities in the Pacific Ocean. Deep-Sea Research II, 46, 2447–2473. Maldonado, M.T., Hughes, M.P., Rue, E.L. and Wells, M.L. (2002) The effect of Fe and Cu on growth and domoic acid production by Pseudo-nitzschia multiseries and Pseudo-nitzschia australis. Limnology and Oceanography, 47, 515–526. Mantoura, R.F.C. and Woodward, E.M.S. (1983) Conservative behaviour of riverine dissolved organic carbon in the Severn Estuary: chemical and geochemical implications. Geochimica et Cosmochimica Acta, 47, 1293–1309. Maranger, R., Bird, D.F. and Price, N.M. (1998) Iron acquisition by photosynthetic marine phytoplankton from ingested bacteria. Nature, 396, 248–251. Margalef, R. (1978) Life forms of phytoplankton as survival alternatives in an unstable environment. Oceanologica Acta, 1, 493–509. Margulis, L. (1993) Symbiosis in cell evolution: microbial communities in the Archaean and Proterozoic eons, W.H. Freeman and Co., New York. Marshall, S.M. (1973) Respiration and feeding in copepods. Advances in Marine Biology, 11, 57–120. Martin, J.H. (1990) Glacial–interglacial CO2 changes: the iron hypothesis. Paleoceanography, 5, 1–13. Martin, J.H. and Gordon, R.M. (1988). North East Pasific Iron distributions in relation to phytoplankton productivity. Deep-Sea Research, 35, 117–196. Martin, J.H., Gordon, R.M. and Fitzwater, S.E. (1991) The case for iron. Limnology and Oceanography, 36, 1793–1802. Mastala, Z., Herodek, S., Balogh, K.V., Borbely, G., Shafik, H.M. and Voros, L. (1996) Nutrient requirement and growth of a Synechococcus species isolated from Lake Balaton. Internationale Revue der Gesamten Hydrobiologie, 81, 503–512. Matsuda, A., Nishijima, T. and Fukami, K. (1999) Effects of nitrogenous and phosphorus nutrients on the growth of toxic dinoflagellate Alexandrium catenella. Nippon Suisan Gakkaishi, 65, 847–855. Middelburg, J.J. and Nieuwenhuize, J. (2000) Nitrogen uptake by heterotrophic bacteria and phytoplankton in the nitrate-rich Thames estuary. Marine Ecology Progress Series, 203, 13–21. Miller, C.B., Morgan, C.A., Prahl, F.G. and Sparrow, M.A. (1998) Storage lipids of the copepod Calanus finmarchicus from Georges Bank and the Gulf of Maine. Limnology and Oceanography, 43, 488–497. Millero, F.J. (1998) Solubility of Fe(III) in sea water. Earth and Planetary Science Letters, 154, 323–329.

358

BIOGEOCHEMISTRY OF MARINE SYSTEMS

Mills, D.K., Tett, P. and Novarino, G. (1994) The Spring Bloom in the south-western North Sea in 1989. Netherlands Journal of Sea Research, 33, 65–80. Minor, E.C., Boon, J.J., Harvey, H.R. and Mannino, A. (2001) Estuarine organic matter composition as probed by direct temperature-resolved mass spectrometry and traditional geochemical techniques. Geochimica et Cosmochimica Acta, 65, 2819–2834. Monod, J. (1942) Recherches sur la Croissance des Cultures Bactériennes, Herman, Paris. Morel, F.M.M., Hudson, R.J.M. and Price, N.M. (1991) Limitation of productivity by trace metals in the sea. Limnology and Oceanography, 36, 1742–1755. NCBI (2002) Taxonomy Browser (Bacteria and Eukaryota pages) http://www.ncbi.nlm.nih.gov/ Taxonomy/(seen August 2002). Nedwell, D.B., Jickells, T.D., Trimmer, M. and Sanders, R. (1999) Nutrients in estuaries. Advances in Ecological Research, 29, 43–92. Nedwell, D.B., Parkes, R.J., Upton, A.C. and Assinder, D.J. (1993) Seasonal fluxes across the sediment–water interface, and processes within North Sea sediments. Philosophical Transactions of the Royal Society of London Series a Mathematical, Physical and Engineering Sciences, 343, 519–529. Nejstgaard, J.C., Gismervik, I. and Solberg, P.T. (1997) Feeding and reproduction by Calanus finmarchicus, and microzooplankton grazing during mesocosm blooms of diatoms and the coccolithophore Emiliania huxleyi. Marine Ecology Progress Series, 147, 197–217. Nejstgaard, J.C., Hygum, B.H., Naustvoll, L.J. and Bamstedt, U. (2001) Zooplankton growth, diet and reproductive success compared in simultaneous diatom- and flagellate-microzooplankton dominated plankton blooms. Marine Ecology Progress Series, 221, 77–91. Nielsen, A. and Richardson, K. (1990) Chrysochromulina polylepis bloom in Danish, Swedish and Norwegian waters, May–June 1988 – an analysis of extent, effects and causes, in Eutrophication and algal blooms in North Sea coastal zones, the Baltic and adjacent areas: prediction and assessment of preventive actions (eds C. Lancelot, G. Billen and H. Barth), Commission of the European Communities, Brussels, Water Pollution Research Report, 12, pp. 11–25. Nodwell, L.M. and Price, N.M. (2001) Direct use of inorganic colloidal iron by marine mixotrophic phytoplankton. Limnology and Oceanography, 46, 765–777. Officer, C.B. and Ryther, J.H. (1980) The possible importance of silicon in marine eutrophication. Marine Ecology Progress Series, 3, 89–91. Ozturk, M., Steinnes, E. and Sakshaug, E. (2002) Iron speciation in the Trondheim Fjord from the perspective of iron limitation for phytoplankton. Estuarine, Coastal and Shelf Science, 55, 197–212. Paasche, E. (1973) Silicon and the ecology of marine plankton diatoms. I. Thalassiosire psuedonana (Cyclotella nana) grown in a chemostat with silicate as a limiting nutrient. Marine Biology, 19, 117–126. Parsons, T.R. and LeBrasseur, R.J. (1970) The availability of food to different trophic levels in the marine food chain, in Marine Food Chains (eds J.H. Steele, Oliver and Boyd), Edinburgh, pp. 325–343. Parsons, T.R., Takahashi, M. and Hargraves, B. (1984) Biological Oceanographic Processes, 3rd edn, Pergamon, Oxford. Pätsch, J. and Radach, G. (1997) Long-term simulation of the eutrophication of the North Sea: temporal development of nutrients, chlorophyll and primary production in comparison to observations. Journal of Sea Research, 38, 275–310. Patterson, D.J. (1999) The diversity of eukaryotes. American Naturalist, 154 (suppl.), S96–S124. Patterson, D.J. and Sogin, M.L. (2001) Tree of Life: Eukaryotes (Eukaryota) http://tolweb.org/tree/ eukaryotes/accessory/treeoverview.html (seen August 2002). Pertola, S., Koski, M. and Viitasalo, M. (2002) Stoichiometry of mesozooplankton in N- and P-limited areas of the Baltic Sea. Marine Biology, 140, 425–434. Philippart, C.J.M., Cadee, G.C., van Raaphorst, W. and Riegman, R. (2000) Long-term phytoplanktonnutrient interactions in a shallow coastal sea: algal community structure, nutrient budgets, and denitrification potential. Limnology and Oceanography, 45, 131–144.

INFLUENCE OF NUTRIENT BIOGEOCHEMISTRY ON ECOLOGY

359

Pierce, R.W. and Turner, J.T. (1992) Ecology of planktonic ciliates in marine food webs. Reviews in Aquatic Sciences, 6, 139–181. Pingree, R.D., Holligan, P.M. and Mardell, G.T. (1978). The effects of vertical stability on phytoplankton distributions in the summer on the Northwest European Shelf. Deep-Sea Research, 25, 1011–1028. Pingree, R.D., Pugh, P.R., Holligan, P.M. and Forster, G.R. (1975) Summer phytoplankton blooms and red tides along tidal fronts in the approaches to the English Channel. Nature, 258, 672–677. Pingree, R.D., Sinha, B. and Griffiths, C.R. (1999) Seasonality of the European slope current (Goban Spur) and ocean margin exchange. Continental Shelf Research, 19, 929–975. Plaza, E., Bosander, J., Dahlberg, A.G. and Hellstrom, B.G. (1990) Operational experience with phosphorus and nitrogen removal at the Himmerfjarden plant, Sweden. Water Science and Technology, 22, 283–284. Pomeroy, L.R. (1974) The ocean’s food web: a changing paradigm. Biological Science, 24, 242–244. Prandle, D., Hydes, D.J., Jarvis, J. and McManus, J. (1997) The seasonal cycles of temperature, salinity, nutrients and suspended sediment in the southern North Sea in 1988 and 1989. Estuarine, Coastal and Shelf Science, 45, 669–680. Prastka, K., Sanders, R. and Jickells, T. (1998) Has the role of estuaries as sources or sinks of dissolved inorganic phosphorus changed over time? Results of a Kd study. Marine Pollution Bulletin, 36, 718–728. Radach, G. (1980) Preliminary simulations of the phytoplankton and phosphate dynamics during FLEX’76 with a simple two-component model. ‘Meteor’ Forschungen-Ergebnisse, A22, 151–163. Radach, G. (1992) Ecosystem functioning in the German Bight under continental nutrient inputs by rivers. Estuaries, 15, 477–496. Radach, G. and Gekeler, J. (1996) Annual cycles of horizontal distributions of temperature and salinity, and of concentrations of nutrients, suspended particulate matter and chlorophyll on the Northwest European shelf. Deutsche Hydrographische Zeitschrift, 48, 261–297. Radach, G., Berg, J. and Hagmeier, E. (1990) Long-term changes of the annual cycles of meteorological, hydrographic, nutrient and phytoplankton time series at helgoland and at LV ELBE 1 in the German Bight. Continental Shelf Research, 10, 305–328. Raven, J.A. (1988) The iron and molybdenum use efficiencies of plant growth with different energy, carbon and nitrogen sources. New Phytologist, 109, 279–287. Raven, J.A. (1990) Predictions of Mn and Fe use efficiencies of phototrophic growth as a function of light availability for growth and C assimilation pathway. New Phytologist, 116, 11–18. Raven, J.A. (1998) The twelth Tansely lecture. Small is beautiful: the picophytoplankton. Functional Ecology, 12, 503–513. Redfield, A.C. (1934) On the proportions of organic derivatives in sea water and their relation to the composition of plankton. James Johnson memorial volume, Liverpool University, Liverpool, UK, 176–192. Redfield, A.C. (1958) The biological control of chemical factors in the environment. American Scientist, 46, 205–221. Redfield, A.C., Ketchum, B.H. and Richards, F.A. (1963) The influence of organisms on the composition of sea water, in The Sea, vol. 2 (ed. M.N. Hill), Interscience, New York, pp. 26–77. Reid, P.C., Borges, M.D. and Svendsen, E. (2001a) A regime shift in the North Sea circa 1988 linked to changes in the North Sea horse mackerel fishery. Fisheries Research, 50, 163–171. Reid, P.C., Holliday, N.P. and Smyth, T.J. (2001b) Pulses in the eastern margin current and warmer water off the north west European shelf linked to North Sea ecosystem changes. Marine Ecology Progress Series, 215, 283–287. Reid, P.C., Lancelot, C., Gieskes, W.W.C., Hagmeier, E. and Weichart, G. (1990) Phytoplankton of the North Sea and its dynamics – a review. Netherlands Journal of Sea Research, 26, 295–331. Rendell, A.R., Ottley, C.J., Jickells, T.D. and Harrison, R.M. (1993) The atmospheric input of nitrogen to the North Sea. Tellus, 45b, 53–63.

360

BIOGEOCHEMISTRY OF MARINE SYSTEMS

Reynolds, C.S. (1987) Community organisation in the freshwater phytoplankton. Symposia of the British Ecological Society, 27, 297–325. Reynolds, C.S. (1996) Plant life of the pelagic. Proceedings of the International Association for Theoretical and Applied Limnology, 26, 97–113. Reynolds, C.S., Irish, A.E. and Elliott, J.A. (2001) The ecological basis for simulating phytoplankton responses to environmental change (PROTECH). Ecological Modelling, 140, 271–291. Riegman, R. (1998) Species composition of harmful algal blooms in relation to macronutrient dynamics, in Physiological Ecology of Harmful Algal Blooms, vol. 41 (eds D.M. Anderson, A.D. Cembella and G.M. Hallegraef), Springer-Verlag, Berlin, Heidelberg, NATO ASI Series G, pp. 475–488. Riegman, R. and Mur, L.R. (1984) Regulation of phosphate uptake kinetics in Oscillatoria agardhii. Archiv für Microbiologie, pp. 28–32. Riegman, R., de Boer, M. and Domis, L.D. (1996) Growth of harmful marine algae in multispecies cultures. Journal of Plankton Research, 18, 1851–1866. Rivkin, R.B. and Smith, E. (1985) Phosphorus metabolism of oceanic dinoflagellates: phosphate uptake, chemical composition and growth of Pyrocystis noctiluca. Marine Biology, 88, 189–198. Roberts, E.C., Davidson, K. and Gilpin, L.C. Response of microplankton communities to N:Si ratio perturbation. Journal of Plankton Research (submitted). Rodrigues, R.M.N.V. and Williams, P.J.L. (2001) Heterotrophic bacterial utilization of nitrogenous and nonnitrogenous substrates, determined from ammonia and oxygen fluxes. Limnology and Oceanography, 46, 1675–1683. Rousseau, V., Leynaert, A., Daoud, N. and Lancelot, C. (2002) Diatom succession, silicification and silicic acid availability in Belgian coastal waters (Southern North Sea). Marine Ecology Progress Series, 236, 61–73. Ryther, J.H. and Officer, C.B. (1981) Impact of nutrient enrichment on water uses, in Estuaries and Nutrients (eds B.J. Nielsen and L.E. Cronin), Humana Press, Clifton, New jersey, pp. 247–261. Sanders, R., Jickells, T. and Mills, D. (2001) Nutrients and chlorophyll at two sites in the Thames plume and southern North Sea. Journal of Sea Research, 46, 13–28. Sanders, R.W. and Wickham, S.A. (1993) Planktonic protozoa and metazoa: predation, food quality and population control. Marine Microbial Food Webs, 7, 197–223. Sandgren, C.D., Hall, S.A. and Barlow, S.B. (1996) Siliceous scale production in chrysophyte and synurophyte algae. 1. Effects of silica-limited growth on cell silica content, scale morphology, and the construction of the scale layer of Synura petersenii. Journal of Phycology, 32, 675–692. Sanudo-Wilhelmy, S.A., Kustka, A.B., Gobler, C.J., Hutchins, D.A., Yang, M., Lwiza, K., Burns, J., Capone, D.G., Raven, J.A. and Carpenter, E.J. (2001) Phosphorus limitation of nitrogen fixation by Trichodesmium in the central Atlantic Ocean. Nature, 411, 66–69. Sciandra, A., Lazzara, L., Claustre, H. and Babin, M. (2000) Responses of growth rate, pigment composition and optical properties of Cryptomonas sp. to light and nitrogen stresses. Marine Ecology Progress Series, 201, 107–120. Seitzinger, S.P. and Giblin, A.E. (1996) Estimating denitrification in North Atlantic continental shelf sediments. Biogeochemistry, 35, 235–260. Seitzinger, S.P. and Sanders, R.W. (1997) Contribution of dissolved organic nitrogen from rivers to estuarine eutrophication. Marine Ecology Progress Series, 159, 1–12. Shaw, P.J., Purdie, D.A., De Freitas, P.S., Ress, D.P. and Joint, I. (1998) Nutrient uptake in a highly turbid estuary (the Humber, United Kingdom) and adjacent coastal waters. Estuaries, 21, 507–517. Sheldon, R.W., Sutcliffe, W.H. and Paranjape, M.A. (1977) Structure of pelagic food chain and relationship between plankton and fish production. Journal of the Fisheries Research Board of Canada, 34, 2344–2353. Simpson, J.H. (1981) The shelf-sea fronts: implications of their existence and behaviour. Circulation and fronts in continental shelf seas (eds R.I. Currie, A.E. Sill, J.H. Simpson and J.C. Swallow), Royal Society, London, 19–31.

INFLUENCE OF NUTRIENT BIOGEOCHEMISTRY ON ECOLOGY

361

Simpson, J.H. and Rippeth, T.P. (1998) Non-conservative nutrient fluxes from budgets for the Irish Sea. Estuarine, Coastal and Shelf Science, 47, 707–714. Smalley, G.W. and Coats, D.W. (2002) Ecology of the red-tide dinoflagellate Ceratium furca: Distribution, mixotrophy, and grazing impact on ciliate populations of Chesapeake Bay. Journal of Eukaryotic Microbiology, 49, 63–73. Smayda, T.J. and Reynolds, C.S. (2001) Community assembly in marine phytoplankton: application of recent models to harmful dinoflagellate blooms. Journal of Plankton Research, 23, 447–461. Smetacek, V.S. (1986) Impact of freshwater discharges on production and transfer of materials in the marine environment, in The role of freshwater outflow in coastal marine ecosystems, vol. G7 (ed. S. Skreslet), Springer-Verlag, Berlin, NATO ASI Series, pp. 85–106. Smith, S.V., Boudreau, P.R. and Ruardij, P. (1997) NP Budget for the Southern North Sea. http:// data.ecology.su.se/mnode/Europe/NorthSea/NORTHSEA.HTM. Sommer, U. (1991) A comparison of the Droop and the Monod models of nutrient limited growth applied to natural-populations of phytoplankton. Functional Ecology, 5, 535–544. Sommer, U. (1994) Are marine diatoms favored by high Si-N ratios? Marine Ecology Progress Series, 115, 309–315. Sondergaard, M., Williams, P.J.L., Cauwet, G., Riemann, B., Robinson, C., Terzic, S., Woodward, E.M.S. and Worm, J. (2000) Net accumulation and flux of dissolved organic carbon and dissolved organic nitrogen in marine plankton communities. Limnology and Oceanography, 45, 1097–1111. Strzepek, R.F. and Price, N.M. (2000) Influence of irradiance and temperature on the iron content of the marine diatom Thalassiosira weissflogii (Bacillariophyceae). Marine Ecology Progress Series, 206, 107–117. Sunda, W. and Huntsman, S. (1997) Interrelated influence of iron, light and cell size on marine phytoplankton growth. Nature, 390, 389–392. Sunda, W.G. (2001) Bioavailability and bioaccumulation of iron in the sea, in The Biogeochemistry of iron in seawater (eds D.R. Turner and K.A. Hunter), pp. 41–84. Sunda, W.G. and Huntsman, S.A. (1995) Iron uptake and growth limitation in oceanic and coastal phytoplankton. Marine Chemistry, 50, 189–206. Sunda, W.G., Swift, D.G. and Huntsman, S.A. (1991) Low Iron requirement for growth in oceanic phytoplankton. Nature, 351, 55–57. Suzumura, M., Ishikawa, K. and Ogawa, H. (1998) Characterization of dissolved organic phosphorus in coastal seawater using ultrafiltration and phosphohydrolytic enzymes. Limnology and Oceanography, 43, 1553–1564. Svensen, C., Egge, J.K. and Stiansen, J.E. (2001) Can silicate and turbulence regulate the vertical flux of biogenic matter? A mesocosm study. Marine Ecology Progress Series, 217, 67–80. Terry, K.L., Laws, E.A. and Burns, D.J. (1985) Growth rate variations in the N:P requirement ratio of phytoplankton. Journal of Phycology, 21, 323–329. Tett, P. (1990) The Photic Zone, in Light and Life in the Sea (eds P.J. Herring, A.K. Campbell, M. Whitfield and L. Maddock), Cambridge University Press, Cambridge, UK, pp. 59–87. Tett, P. (1992) The ecology of plankton in Scottish coastal waters. Proceedings of the Royal Society of Edinburgh, 100B, 27–54. Tett, P. (1998) Parameterising a microplankton model. Napier University, Edinburgh, Report, ISBN 0 902703 60 9. Tett, P. and Droop, M.R. (1988) Cell quota models and planktonic primary production, in Handbook of Laboratory Model Systems for Microbial Ecosystems, vol. 2 (ed. J.W.T. Wimpenny), CRC Press, Florida, pp. 177–233. Tett, P. and Edwards, V. (2003) Review of Harmful Algal Blooms in Scottish coastal waters. Scottish Environmental Protection Agency, Stirling, Report. Tett, P. and Mills, D. (1991) The plankton of the North Sea – pelagic ecosystems under stress? Ocean and Shoreline Management, 16, 233–257.

362

BIOGEOCHEMISTRY OF MARINE SYSTEMS

Tett, P. and Walne, A. (1995) Observations and simulations of hydrography, nutrients and plankton in the southern North Sea. Ophelia, 42, 371–416. Tett, P., Drysdale, M. and Shaw, J. (1981) Phytoplankton in Loch Creran in 1979, and its effect on the rearing of oyster larvae. Scottish Marine Biological Association, Oban, Internal report, 52, 77 pp. Tett, P., Gilpin, L., Svendsen, H., Erlandsson, C.P., Larsson, U., Kratzer, S., Fouilland, E., Janzen, C., Lee, J.-Y., Grenz, C., Newton, A., Ferreira, J.G., Fernandes, T. and Scory, S. Eutrophication and some European waters of restricted exchange. Nearshore and Coastal Oceanography (accepted). Tett, P., Gowen, R., Grantham, B., Jones, K. and Miller, B.S. (1986) The phytoplankton ecology of the Firth of Clyde sea-lochs Striven and Fyne. Proceedings of the Royal Society of Edinburgh, 90B, 223–238. Tett, P., Heaney, S.I. and Droop, M.R. (1985) The Redfield ratio and phytoplankton growth rate. Journal of the Marine Biological Association of the United Kingdom, 65, 487–504. Tett, P., Joint, I., Purdie, D., Baars, M., Oosterhuis, S., Daneri, G., Hannah, F., Mills, D.K., Plummer, D., Pomroy, A., Walne, A.W. and Witte, H.J. (1993) Biological consequences of tidal stirring gradients in the North Sea. Philosophical Transactions of the Royal Society of London Series a Mathematical, Physical and Engineering Sciences, 340, 493–508. Tett, P., Kennaway, G.M., Boon, D., Mills, D.K., O’Connor, G.T., Walne, A.W. and Wilton, R. (2001) Optical monitoring of phytoplankton blooms in Loch Striven, a eutrophic fjord. International Journal of Remote Sensing, 22, 339–358. Tilman, D., Kilham, S.S. and Kilham, H. (1982) Phytoplankton community ecology – the role of limiting nutrients. Annual Review of Ecology and Systematics, 13, 349–372. Timmermans, K.L., Stolte, W. and Debaar, H.J.W. (1998) Responses of marine phytoplankton in iron enrichment experiments in the northern North Sea and north east Atlantic Ocean. Marine Chemistry, 61, 229–242. Timmermans, K.R., Stolte, W. and Debaar, H.W. (1994) Iron-mediated effects on nitrate reductase in marine phytoplankton. Marine Biology, 121, 389–396. Tomas, C.R. (ed.) (1997) Identifying marine phytoplankton, San Diego & London, Academic Press, xv + 858. Trimmer, M., Gowen, R.J., Stewart, B.M. and Nedwell, D.B. (1999) The spring bloom and its impact on benthic mineralisation rates in western Irish Sea sediments. Marine Ecology Progress Series, 185, 37–46. Turner, D.R. and Hunter, K.A. (eds) (2001) The Biogeochemistry of iron in seawater, John Wiley & Sons, Chichester. Vadstein, O. (1994) The role of heterotrophic, planktonic bacteria in the cycling of phosphorus in lakes: phosphorus requirements, competitive ability and food web interactions. PhD thesis, University of Trondheim (cited by Gismervik et al., 1996). Verity, P.G. (2000) Grazing experiments and model simulations of the role of zooplankton in Phaeocystis food webs. Journal of Sea Research, 43, 317–343. Verity, P.G. and Paffenhofer, G.A. (1996) On assessment of prey ingestion by copepods. Journal of Plankton Research, 18, 1767–1779. Verity, P.G. and Smetacek, V. (1996) Organism life cycles, predation, and the structure of marine pelagic ecosystems. Marine Ecology Progress Series, 130, 277–293. Verity, P.G., Williams, S.C. and Hong, Y. (2000) Formation, degradation, and mass:volume ratios of detritus derived from decaying phytoplankton. Marine Ecology Progress Series, 207, 53–68. Visser, M., Batten, S., Becker, G., Bot, P., Collin, F., Damm, P., Danielssen, D., van den Eynde, D., Foeyn, L., Frohse, A., Groeneveld, G., Laane, R. and van Raaphorst, W. (1996) Time series analysis of monthly mean data of temperature, salinity, nutrients, suspended matter, phyto- and zooplankton at eight locations on the Northwest European Shelf. Deutsche Hydrographische Zeitschrift, 48, 299–323. de Vries, I., Duin, R.N.M., Peeters, J.C.H., Los, F.J., Bokhorst, M. and Laane, R.W.P.M. (1998) Patterns and trends in nutrients and phytoplankton in Dutch coastal waters: comparison of time-

INFLUENCE OF NUTRIENT BIOGEOCHEMISTRY ON ECOLOGY

363

series analysis, ecological model simulation, and mesocosm experiments. ICES Journal of Marine Science, 55, 620–634. Wassmann, P., Egge, J.K., Reigstad, M. and Aksnes, D.L. (1996) Influence of dissolved silicate on vertical flux of particulate biogenic matter. Marine Pollution Bulletin, 33, 10–21. Wheeler, D.L., Chappey, C., Lash, A.E., Leipe, D.D., Madden, T.L., Schuler, G.D., Tatusova, T.A. and Rapp, B.A. (2000) Database resources of the National Center for Biotechnology Information. Nucleic Acids Research, 28, 10–14. Whittaker, R.H. (1969) New concepts of kingdoms of organisms. Science, 163, 150–160. Widdicombe, C.E., Archer, S.D., Burkill, P.H. and Widdicombe, S. (2002) Diversity and structure of the microplankton community during a coccolithophore bloom in the stratified northern North Sea. Deep-Sea Research II – Topical Studies in Oceanography, 49, 2887–2903. Williams, P.J.L. (1981) Incorporation of microheterotrophic processes into the classical paradigm of the plankton food web. Kieler Meeresforschungen, 5, 1–28. Williams, P.J.L. (1995) Evidence for the seasonal accumulation of carbon-rich dissolved organic material, its scale in comparison with changes in particulate material and the consequential effect on net C/N assimilation ratios. Marine Chemistry, 51, 17–29. Williams, P.J.L. and Egge, J.K. (1998) The management and behaviour of the mesocosms. Estuarine, Coastal and Shelf Science, 46, (supl. A) 3–14. Woese, C.R., Kandler, O. and Wheelis, M.L. (1990) Owards a natural system of organisms – proposal for the domains Archaea, Bacteria, and Eucarya. Proceedings of the National Academy of Sciences of the United States of America, 87, 4576–4579. Wu, J. and Luther, G.W. (1995) Complexation of Fe(III) by natural organic ligands in the north west Atlantic ocean, as determined by competitive equilibration method and kinetic approach. Marine Chemistry, 50, 159–177. Yamaguchi, M. and Itakura, S. (1999) Nutrition and growth kinetics in nitrogen- or phosphorus-limited cultures of the noxious red tide dinoflagellate Gymnodinium mikimotoi. Fisheries Science, 65, 367–373. Yamaguchi, M., Itakura, S. and Uchida, T. (2001) Nutrition and growth kinetics in nitrogen- or phosphorus-limited cultures of the ‘novel red tide’ dinoflagellate Heterocapsa circularisquama (Dinophyceae). Phycologia, 40, 313–318.

Index

ABE 279 accretionary prisms 267 acid volatile sulphide 21, 71 acidifying gases 143 aerial roots 31 aerobic respiration 15 Alaska 145 Aleutian Trench 270 Alexandrium minutum 314 algal blooms 68 alien species 147 alkaline phosphatase 50 alveolata 304 Amazon river 243 ammonium 48 fluxes 17 anhydrite 242 anoxic conditions 165 reef pore-waters 53 waters 76 anthropogenic impacts on shallow reef 57 anthropogenic perturbations 276 anti-cyclonic circulation regime 139 anti-estuarine 91, 92, 93 aquaculture industry 68 ponds 1 Arabian Sea 157 Archaea 254, 303 Archaean cherts 260 Arctic 127 marginal seas 129 ocean 127 oscillation 139 shelves 133 aromatic hydrocarbons 268 assimilatory nitrate reduction 186 Asterionella 305 Aswan Dam 96 Atlantic waters 132 atmospheric carbon dioxide 224 atmospheric deposition 297 attenuation coefficient 172 Aureococcus 305

Aureoumbra 305 lagunensis 311 authigenic enrichments 85 autonomous vehicles 279 autotrophs 308 Axial Volcano 264 Bacillariophyceae 305 Bacillariophyta 305 back-arc spreading centres 241 back-reef 41 bacterial abundance 175 production 174 weathering 260 balance of organisms 294 ballast water 147 Baltic Sea 326, 340 Barents Sea 128, 136 barite 242, 269 basaltic glass 266 bathymodiolid mussels 240 Bay of Bengal 160 beam attenuation 171 Beggiatoa 268 benthic algae 8 carbon remineralisation 197 communities 195 nutrient pools 2 organisms 28 processes 195 benthic–pelagic coupling 137 benthos 137, 138, 228 Bering Strait 128 Bicoesida 305 bioamplification 144 biodiversity 138, 268 biofilms 255 biogeography 268 biomarkers 134 biomass 239 biome 345 biomineralisation 256 bioproduction 136

INDEX Biosphere 2 coral reef 45, 55 bioturbation 2, 20, 216, 218 Black Sea 116 black smokers 241 blooms 163 Bodo 304 bottom-up factors 312 brine 128 C:N ratio 319 C:N:P ratios 55 C:P ratio 46, 51 Cafeteria 305 Calanus helgolandicus 313 Canada 146 Canadian Archipelago 128 Basin 128 carbon cycle 263 cycling at vents 252 deposition 47 dioxide 185 dioxide flux to atmosphere 185 mineralization 12 reef 43, 44 carbonate–brucite chimneys 245 carbonates, microbial 274 carbonic acid 52 carotenoid 311 cell-quota theory 320 CEPEX 332 Ceratium 304, 310 Ceratium furca 312 Chaetoceros 305 chalcopyrite 242 Chatonella 305 chelation, organic 302 chemical tracers 76 chemocline 80 chemosynthetic 239 symbionts 267 chimney 254 Chlorococcales 305 Chlorophyceae 305 chlorophyll 107 Chlorophyta 305 Chloroxybacteria 304 choanoflagellates 305 chromium reducible sulphide 21 Chroococcales 304 Chrysochromulina 305, 309, 314 Chrysophyta 305

365

Chukchi Sea 128 Chytridomycota 305 chytrids 305 Ciliates 304 Ciliophora 304 cladoceran crustaceans 314 clams, Calyptogena 270 Solemya 270 climate change 138 feedback 278 models 141 shifts 139 variability 138 warming 141 coccolithophores 181 Coccoshpaerales 305 cod stocks 139 cold-condensation effect 144 cold-core eddy 96, 98 commercial fishing 145 shipping route 146 community production 46 respiratory quotients CRQ 15 competition experiments 337 competitive exclusion principle 307 contaminants 143 copepods 193, 313, 314 copper stress 314 co-precipitation 75 Corethron 169 Coscinodiscophyceae 305 Coscinodiscus 170, 310 coupled nitrification–denitrification 18 crabs 16 crude oil 268 cryptomonad 304, 311 Cryptomonas 304 Cryptophyta 304 CSR conceptual model 307 cyanobacteria 102, 304, 307, 311 Cyanophyceae 304 cycling of carbon 77 cyclonic circulation regime 139 CYPRIS station 300 Cyprus eddy 98 DAIN 294 decomposition constants 11 deep chlorophyll maximum (DCM) 91, 98, 100 denitrification 16, 50, 135, 165, 186, 299, 300 sedimentary 195

366 Desulfovibrio 272 detritus 7 diagenesis 69 diatom-copepod-fish trophic paradigm 313 diatoms 111, 170, 305 centric 307 pennate 307 tychopelagic 307 Dictyocha speculum 320 Dictyochophyceae 305 diffusive boundary layer 49, 54 DIN 166 Dinobryon 305 dinoflagellates 111, 304, 307, 314 Dinophyceae 304 Dinophysidae 304 Dinophysis 304 Dinophyta 304 Dissolved Available Inorganic Nitrogen 294 dissolved inorganic carbon 223 organic carbon 5, 47, 175 organic matter 4 organic nitrogen 5, 100, 298 organic phosphorus 100 organic pools 295 distribution coefficient 295 divergent plate boundaries 241 diversity 239, 293 DNA in vent fluids 246 doliolid salps 314 domoic acid 314 downwelling 166 DROOP model 320 East Greenland Current 131 ecological economics 144 ecosystem switch 187 eddies 160 Eel River Basin 273 El Niño–Southern Oscillation 140, 221, 222 electron transport system 187 element cycling 66 elemental sulphur 72 Emiliana 305 huxleyi 312 ENCORE experiments 57 environmental changes 138 enzymes, heat stable 251 eolian supply of dust 198 EPR 258

INDEX erosive margins 267 ERSEM 331 estuarine 296 Eubacteria 254, 303 Eucarya 303 eucaryotes 102 Euglenida 304 Eugleniphyceae 304 Euglenozoa 304 euphotic zone 102 Eurasian Basin 128 Eustigmatophyta 305 Eutreptiella 304 eutrophic equatorial abyss 214 event plumes 244 fast ice 132 fecal pellets 27, 184 feedback loops 138 fertilizer inputs from land 189 filaments 160 filtering effect 296 fish 140 processing 145 flood 75 floristic composition 307, 308, 310 flushing events 67 food web structure 179 foraminifera 181, 304 fore-reef 41 forest age 25 type 25 formaldehyde 51 fossil fuels 278 fossilised tube structures 259 Fragilariophyceae 305 Fram Srait 128 Framvaren Fjord 77 f-ratios 167 Fridtjof Nansen 140 galena 242 Gallionella 257 gas emission 185 hydrates 268 geological stability 276 global warming 141, 223 potential 278 Gonyaulacales 304 Gonyaulax 304

INDEX Gorda Ridge 259 greenhouse effect 132 Greenland 144, 146 Sea 131 groundwater discharge 297 Guaymas Basin 256, 274 Gulf of Mexico 270 Gymnodiniales 304 gyres 98 Gyrodinium 304 haptophytes 305, 307, 312 harbour seal 140 hazardous chemicals 143 heavy metals 143 Helgoland 329 herbivorous crabs 26 herring stocks 139 Heterokonts 305 Heterosigma 305 heterotrophic 239 bacteria 104, 174, 319 heterotrophic dinoflagellates 313, 319 heterotrophs 308 high carbon production 40 high nitrate low chlorophyll 345 high turbidity 5 Himmerfjord 337 hull fouling 147 hunters 145 Hydrate Ridge 273 hydraulic conductivity 53 hydrocarbon fluxes 281 hydrodynamics 3 hydrogen 264 oxidising bacteria 252 sulphide 239 hydrothermal fluids 242 heat flux 245 plumes 245, 252 hydrothermophilic microbes 246 ice algae 136 edge 131 fauna 136 formation 132 melting 132 ice-associated community 132 iceberg scour 138 Iheya Basin 258

367

impacts on coral reefs 58 incubation chambers 58 Indian Ocean 157 indigenous peoples 128, 144 Indo-Pacific reef flats 57 intermediate nepheloid layer 191 intraplate volcanism 244 iodate 51 iodine 51, 192 Irish Sea 296, 299, 300, 340 iron 192 associated with particles 302 colloidal 327 concentration, variations between oceans 301 deficiency 314 dissolved 301 enrichment 326 fertilisation 229 forms of 302 hydroxides 6 limitation 319, 345 limited 117 oxide muds 257 oxidisers 252 reduction 19 uptake 302 iron:phosphate ratio 323 iron-bound phosphate 24 iron-oxidising bacteria 261 Isochrysidales 305 isotope enrichments 78 biomarkers 281 fractionation 78 Japan Trench 267 Juan de Fuca Ridge 245 Kd 295 kelp forest 140 Kelvin waves 159 killer whale 140 K-selected 310 Labrador Sea 131, 139 laminated sediments 196 landslide 75 Lau basin 263 layer-silicates 258 leads 136 leaf-eating crabs 9

368 Leptocylindrus 305 Leptospirillum 257 Leptothrix 257 Levantine deep water 94 Levantine intermediate water 91 lifeforms 307 light limited 5 light-nutrient-mixing theory 309 Lingulodinium machaerophorum 310 lipid 136, 143 litter-bags 9 litter fall 7 Loch Creran 340 Loch Striven 340 LOICZ 299 Loihi Seamount 257 Lost City hydrothermal field 245 macrofauna 196 macrophyte production 8 magmatic degassing 241 management 146 manganese 192 nodules 218, 226 manganese-oxidising bacteria 252 mangrove crabs 7 creek 3 peats 23 mantle-derived methane 251 Manus basin 263 marcasite 242 marginal ice zone 135 marine mammals 145 mass flux 243 transfer relationships 55 MECHANISTIC models 325 Mediterranean outflow water 95 megafauna 228 Mendocino Fracture Zone 267 mercuric chloride 51 mercury pollution 73 Mersah Matru eddy 98 mesocosm experiments 332, 333 Mesodinium 304 mesophiles 251 mesoscale eddies 94 mesotrophic sub-equatorial abyss 217 mesozooplankton 177 metal mobility 240 metaliferous sediments 245 Metallogenium 257

INDEX metals, hydrothermal contribution to seawater 245 methane 132, 239 oxidation 83, 240, 272, 273 plumes 269 methane-producing bacteria 23 methanogenesis 239, 271 methanotrophs 272 methyl mercury 73 microbial layer 80 loop 184, 313 mineralisation 6 uptake 81 micro-organism taxonomy 303 microplankton 303 microzooplankton 175 grazing 180 Mid-Atlantic ridge 245 Middle Valley 256 mid-ocean-ridge 241 mining 277 mixed layer dynamics 172 layers 162 model, plankton 308 molecular phylogeny 240 MONOD theory 320 monsoon 2, 159 northeast 159 southwest 159 mother populations 277 mud volcanoes 267 myctophids 192 Myrionecta 304 myxotrophs 308 N fixing phytoplankton 118 N:P ratio 5, 53, 55, 311, 344 winter 301 N:Si ratio 332, 342 Nannochloropsis 305 native groups 145 natural capital 144 NeMO 279 net ecosystem production 299 neuston 111 new production 166 N-fixation 166 Nile 108 nitrate 48, 98 deficits 187 fluxes 18 uptake 183

INDEX nitrification 18, 50, 186 nitrite 48 reductase 319 nitrogen budget 298 fixation 50 oxidation states 294 reef 44 nitrogenase 326 Nitromonas 304 nitrous oxide 188 Nitzschia 170 Noctiluca 304 Noctilucales 304 nodule mining 226, 227 NOEWESP project 300 non-motile greens 307 nontronite 258 North Atlantic 165 net nitrogen flux 299 Oscillation 139 North Sea 296, 297, 312, 329 Project 297 Norway 146 Norwegian Sea 136 Nostocales 304 nutricline 97 nutrient concentrations 98 cycling 4 element ratios 319, 328 levels 4 limitation 116 loading of coral reefs 56 trap 164 uptake 46 nutrients 163 budget for North Sea 297 in coral reef waters 48 dissolved organic 312 sink 298 sorption to inorganic particles 295 oceanic biological pump 177 ochre 257 Ochromonas 305, 312 Ochromonodales 305 ODP 256 off-axis 243 oil exploration 277 Okinawa Trough 258 oligotrich ciliates 313 oligotrophic central gyre abyss 218 opal 181

ophiolites 243 Opsithokonts 305 Oregon margin 270 organic carbon accumulation 195 detritus 298 matter decomposition 16 matter preservation 85 nitrogen 48 Oscillatoriales 304 oxidation of H2S 82 oxides 244 oxygen conditions 67 consumption, sediment community 197 depletion 165 penetration 15 uptake 12, 13, 14, 15 oxygen-deficient zones 185 oxyhydroxides 242 Oxyrrhinaceae 304 P limited 116 Pacific Decadal Oscillation 221, 222 Pacific waters 132 pack ice 136 paradox of the plankton 307 Paraphysomonadaceae 305 particle aggregation 194 particle flux 180, 263 particulate organic matter 295 Pavlova 305 Pavlovales 305 PCB 69 pelagic heterotrophic community 6 tunicates 314 Pelagophyceae 305 Peridiniales 304 Peridinium trochoideum 310 peridotite 245 permanent ice pack 132 Persian Gulf 163 persistent organic contaminants 143 Peru margin 267, 270 petroleum 146, 268 pH changes 19 Phaeocystis 170, 305, 307, 314 Phagotrophic myxotrophs 312 phagotrophs 308 phosphate 98 authigenic 196 cycling 23 flux 24, 45

369

370 phosphorus enrichment 74 limited 91, 105 limiting 5 reef 44, 50 requirement 294 photoredox cycling 302 photosynthesis 45, 239 phycobilins 311 phycobiliprotein 304 physical disturbance 138 phytoplankton 8, 111 bio-optical properties 173 bloom 114, 173 community structure 167 composition 167 growth 180 growth and mortality 179 size distribution 167 pico-plankton 167 pigment composition 86 pigments 197 Pleuromamma indica 179 POC 163, 263 flux 211, 213, 220, 221, 225 polymetallic massive sulphides 277 polynyas 133 POM 295 pore water 72 pools 10 residence time 53 Porosira 169 Prasinophyceae 305 prawn farming 31 prey size 314 primary nitrite maximum 186 production 5, 105, 163, 299 productivity 97, 253 Prochlorococcus 169, 304, 311 prochlorophytes 102 Prorocentrales 304 Prorocentrum 304 micans 310 protected areas 278 Proteobacteria 304 Protoctista 303 Protoperidinium 304 prymeiophytes 104 Prymnesiales 305 Pseudomonas 304 Pseudo-nitzchia 305 Pseudo-nitzschia multiseries 314

INDEX pulp and paper industries 70 pyrite 71, 242 pyrite formation 82 Pyrocystales 304 Radiolaria 304 radionuclides 143 Rainbow MAR 263 Raphidophyta 305 rare earth element 78 reactive iron 72 recycled nutrients 329 Red Sea 163 red tide dinoflagellates 323 Redfield 315 Redfield ratio 6, 133, 293 flexible 347 redox 11, 213 potential 254 reactions of nitrogen 49 redoxcline 81 redox-sensitive elements 192 reducing conditions 189 reduction in iodate 192 reef crest 41 flat 41 morphology 41 zonation 41 reforestation 30 regime shifts 138 remineralisation 298 residence times 3 resilience 277 resource extraction 277 respiration, reef 45 respiratory quotient 45 reverse methanogenesis 83 Rhizosolenia 169, 310 Rhodes Gyre 95, 98 ridge flanks 243 River Po 115 river runoff 159 riverine inputs of nutrients 299 Rossby waves 159 r-selected 310 Russia 146 salps 184 salt tectonics 267 sapropels 92 saprotrophs 308

INDEX Sarcodina 304 Scandinavia 144 scavenging 73 Sea Cliff hydrothermal field 259 sea ice 127 lion 140 otter 140 urchin 140 seabirds 140 seafloor observatories 279 seagrass 29 leaves 11 seamounts 241 seasonal thermocline 100 variations 28 seawater-basalt interaction 242 SeaWiFS 113 sediment structure 12 trap 181 –water interface 68 sedimentation 163 rates 65 sediment–water interface 12, 26 Serial Endosymbiosis Theory 303 sesarmide crabs 26 sesarmine crab 2, 12 shear stress 3 shelf break 297 shelf-basin exchange 134 Shikmona Gyre 108 shrimp farming 7, 9, 23 Si/Ca ratio 181 Si:N:P ratio 311 Si:P ratio 311 siderophore 302, 325 silica 242 reef 51 requirement 319 silicate 98 requirement 294 Silicoflagellates 305 sill 67 size-fractionated chl a and primary production 168 Skagerrak 341 Skeletonema 305 slumping 65 Snake Pit 255 snowblower vents 246, 249 solar insolation 4 Southampton water 310

Southern Explorer 258 sphalerite 242 stable carbon isotopes 264 Stanton number 54 Straits of Sicily 94 Stramenophiles 305 stratification 173 stromatolites 275 Strombidium 304 suboxic sediments 76 suboxic zone 189 subsistence harvesting 145 quotas 321 subsurface biosphere 246 chlorophyll maximum 172 Sulfur Springs 260 sulphate reduction 15, 19, 22, 70, 81, 189, 240, 271 sulphate-reducers 251 sulphates 242 sulphide mounds 254 oxidation 240, 272 sulphides 242 sulphur oxidisers 249, 252 sulphur-rich floc 246 surface currents 159 suspended load 5 symbiosis 40 symbiotic 239 bacteria 48 Synechococcus 103, 169, 304, 311 Synura petersenii 320 Synurales 305 TAG 255 Thailand 1 Thalassiosira 305 thermohaline circulation 131 thermophilic organisms 246 Thioploca 196 thiosulphate 265 thiotrophs 272 tidal asymmetry 3 exchange 7 mixing fronts 309 Tintinnopsis 304 top-down trophic effects 312 Transpolar Drift 129 transport time in NW European shelf seas 301 Trichodesmium 166, 304

371

372 trophic behaviour 268 trophy 308 ultra-oligotrophic 91 ultraviolet-B radiation 127 underwater mining 277 United States 146 upwelling 160 urbanisation 9 varved sediments 196 vertical migration 179 vesicomyid clams 240 vestimentiferan worms 240 Vietnam 1 viral biomass 262 Viridioplantae 305 vitamin B12 312

INDEX volatiles 241 volcanic-hosted massive sulphide (VMS) 243 Volvocales 305 Wadden Sea 340 warm-core eddy 98 water, transport through reef framework 52 water-rock reactions 241 wave-induced mixing 53 West Spitsbergen Current 128 Western Mediterranean 93 wind stress 185 winter convection 166, 181 wurtzite 242 zooplankton 111, 264, 303 biomass 177, 179

Jan

Apr

Feb

May

Mar

Jun

July

Oct

Aug

Nov

Sept

Dec

Color Plate 1 SeaWiFS 9-km monthly composite chlorophyll images for 2000; chlorophyll values in mg m−3 shown on the colour scale. SeaWiFS images courtesy of NASA SeaWiFS Project and Orbimage Inc. (see Fig. 4.7).

Chl a (mg/m3)

Temperature (°C)

25 20 15 10 5

Latitude (N)

25 20 15 10 5 25 20 15 10 5 50

0.0

55

0.4

60

0.8

65

70

1.5

75 50 55 Longitude (E) 4.0

18

22

60

65

26

70

30

75

34

Color Plate 2 Monthly climatologies of remotely sensed surface chlorophyll a (a–c) and SST (d–f) for February (a, d), May (b, e) and August (c, f). Chlorophyll and SST climatologies were created by Jerry Wiggert (University of Maryland, College Park) and Bob Evans (RSMAS, University of Miami) using the SeaWiFS (Sea-viewing Wide Field-of-view Sensor) and MODIS (Moderate-resolution Imaging Spectroradiometer) data, respectively. Note that some features of circulation (e.g. the great whirl) can be readily identified in both sets of data (see Fig. 6.2).

Color Plate 3 Distributions of chlorophyll fluorescence (panels on the left) and POC (panels on the right) in the upper 150 m along the US JGOFS southern transect during different seasons. These properties were derived from in situ light transmission and fluorescence measurements. The dotted and solid lines indicate the mixed layer depth (identified by an increase in σθ by 0.03 relative to the surface) and the 0.5 μM NO3− contour, respectively. Reproduced from Gundersen et al. (1998) with permission from Elsevier Science (see Fig. 6.8).

Color Plate 4 Acoustic backscatter for 24 hours around 19° N, 67° E indicating diurnal migration of organisms (mostly myctophids). Reproduced from Morrison et al. (1999) (see Fig. 6.18).

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