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Nitrogen Cycling inCoastal MarineEnvironments
Edited by
T. Henry Blackburn
and
Jan S~rensenDepartment of Ecology and Genetics
University of Aarhus, Denmark
Proceedings from a SCOPE symposium held at the University of Aarhus,3-7 June 1985
Published on behalf of theScientific Committee on Problems of the Environment (SCD PE)
of theInternational Council of Scientific Unions (ICSU)
byJOHN WILEY & SONS
Chichester. New York. Brisbane. Toronto. Singapore
Copyright cg 1988 by theScientific Committee on Problems of the Environment (SCOPE)
All rights reserved.
No part of this book may be reproduced by any means,or transmitted, or translated into a machine language without thewritten permission of the copyright holder.
Library of Congress Cataloging-in-Publication Data:
Nitrogen cycling in coastal marine environments.
(SCOPE; 33)Includes Index,1. Nitrogen cycle. 2. Marine ecology. 3. Coastal
ecology. I. Blackburn, T. Henry (Thomas Henry),1933- . II. S~rensen, Jan Poul. III. InternationalCouncil of Scientific Unions. Scientific Committee onProblems of the Environment. IV. Series: SCOPE(Series); 33.QH344. N54 1988 574.5'2638 86-26803
ISBN 0471 914045
British Library Cataloguing in Publication Data:
Nitrogen cycling in coastal marine environments.(Scope; 33)1. Marine ecology 2. Nitrogen cycleI. Blackburn, T. Henry II. S~rensen, JanIII. International Council of ScientificUnions. Scientific Committee on Problems ofthe EnvironmentIV. Series574.5'2636 QH541.5.S3
ISBN 0471 914045
Phototypeset by Thomson Press (India) Limited, New DelhiPrinted and bound in Great Britain by Anchor Brendon Ltd., Tiptree, Essex.
ToSheila andAnni
SCOPE 1:
SCOPE 2:
SCOPE 3:
SCOPE 4:
Global Environmental Monitoring 1971, 68pp (out of print)
Man-Made Lakes as Modified Ecosystems, 1972, 76pp
Global Environmental Monitoring Systems (GEMS): Action Planfor Phase 1, 1973, 132pp
Environmental Sciences in Developing Countries, 1974, 72pp
Environment and Development, proceedings of SCOPE/UNEP Symposium onEnvironmental Sciences in Developing Countries, Nairobi, February 11-23,1974, 418pp.
SCOPE 5:
SCOPE 6:
SCOPE 7:
SCOPE 8:
SCOPE 9:
SCOPE 10:
SCOPE 11:
SCOPE 12:
Environmental Impact Assessment: Principles and Procedures,Second Edition, 1979, 208pp
Environmental Pollutants: Selected Analytical Methods, 1975,277pp (available from Butterworth & Co. (Publishers) Ltd,Sevenoakes, Kent, England
Nitrogen, Phosphorus, and Sulphur: Global Cycles, 1975, 192pp(available from Dr Thomas Rosswall, Swedish Natural ScienceResearch Council, Stockholm, Sweden)
Risk Assessment of Environmental Hazard, 1978, 132pp
Simulation Modelling of Environmental Problems, 1978, 128pp
Environmental Issues, 1977, 242pp
Shelter Provision in Developing Countries, 1978, 112pp
Principles of Ecotoxicology, 1978, 372pp
SCOPE 13: The Global Carbon Cycle, 1979, 491pp
SCOPE 14: Saharan Dust: Mobilization, Transport, Deposition, 1979, 320pp
SCOPE 15: Environmental Risk Assessment, 1980, 176pp
SCOPE 16: Carbon Cycle Modelling, 1981, 404pp
SCOPE 17: Some Perspectives of the Major Biogeochemical Cycles, 1981,175pp
SCOPE 18: The Role of Fire in Northern Circumpolar Ecosystems, 1983,344pp
SCOPE 19: The Global Biogeochemical Sulphur Cycle, 1983, 495pp
SCOPE 20: Methods for Assessing the Effects of Chemicals on ReproductiveFunctions, 1983, 568pp
VB
Vlll
SCOPE 21: The Major Biogeochemical Cycles and Their Interactions, 1983,554pp
SCOPE 22: Effects of Pollutants at the Ecosystem Level, 1984, 443pp
SCOPE 23: The Role of Terrestrial Vegetation in the Global Carbon Cycle:Measurement by Remote Sensing, 1984, 272pp
SCOPE 24: Noise Pollution, 1986
SCOPE 25: Appraisal of Tests to Predict the Environmental Behaviour ofChemicals, 1985, 400pp
SCOPE 26: Methods for Estimating Risks of Chemical Injury: Human andNon-human Biota, 1985, 712pp
SCOPE 27: Climate Impact Assessment: Studies of the Interaction of Climateand Society, 1985, 649pp
SCOPE 30:
Environmental Consequences of Nuclear WarVolume I Physical, 1985, 342ppVolume II Ecological and Agricultural Effects, 1985, 523pp
The Greenhouse Effect, Climate Change and Ecosystem, 1986,574pp
Methods for Assessing the Effects of Mixtures of Chemicals, 1987,928pp
SCOPE 28:
SCOPE 29:
SCOPE 31: Occurrence and Pathways of Lead, Mercury, Cadmium andArsenic in the Environment, 1987, 384pp
SCOPE 32: Land Transformation in Agriculture, 1987, 580pp
SCOPE 33: Nitrogen Cycling in Coastal Marine Environments, 1988, 488pp
Funds to meet SCOPE expenses are provided by contributions from SCOPENational Committees, an annual subvention from ICSU (and through ICSU,from UNESCO), an annual subvention from the French Ministere del'Environment et du Cadre de Vie, contracts with UN Bodies, particularlyUNEP, and grants from Foundations and industrial enterprises.
International Council of Scientific Unions (ICSU)Scientific Committee on Problems of the Environment (SCOPE)
SCOPE is one of a number of committees established by a non-governmentalgroup of scientific organizations, the International Council of Scientific Unions(ICSU). The membership of ICSU includes representatives from 68 NationalAcademies of Science, 18 International Unions and 12 other bodies calledScientific Associates. To cover multidisciplinary activities which include theinterests of several unions, ICSU has established 10 scientific committees, ofwhich SCOPE is one. Currently, representatives of 34 member countries and15 Unions and Scientific Committees participate in the work of SCOPE, whichdirects particular attention to the needs of developing countries. SCOPE wasestablished in 1969in response to the environmental concerns emerging at thattime; ICSU recognized that many of these concerns required scientific inputsspanning several disciplines and ICSU Unions. SCOPE's first task was toprepare a report on Global Environmental Monitoring (SCOPE 1, 1971)for theUN Stockholm Conference on the Human Environment.
The mandate of SCOPE is to assemble, review, and assess the informationavailable on man-made environmental changes and the effects of these changeson man; to assess and evaluate the methodologies of measurement of environ-mental parameters; to provide an intelligence service on current research; and bythe recruitment of the best available scientific information and constructivethinking to establish itselfas a corpus of informed advice for the benefit of centresof fundamental research and of organizations and agencies operationallyengaged in studies of the environment.
SCOPE is governed by a General Assembly, which meets every three years.Between such meetings its activities are directed by the Executive Committee.
R. E. MunnEditor-in-ChiefSCOPE Publications
Executive Secretary: V. Plocq
Secretariat: 51 Bid de Montmorency75016 PARIS
IX
Acknowledgements
We are indebted to Professor Thomas Rosswall, University of Linkoping,Sweden, who suggested a SCOPE Symposium on Nitrogen Cycling in CoastalMarine Environments be held at our Department and also organized part of thefunding. His encouragement in our preparation of the meeting was most valuable.
Special thanks are due to the participants who contributed with seminars,posters and manuscripts. We also thank our colleagues and students who assistedus with their enthusiasm during the meeting.
Sponsors of the Symposium were the Scientific Committee on Problems of theEnvironment (SCOPE), Danish Natural Science Research Council, and SwedishBoard for Environmental Protection. The financial support is gratefullyacknowledged.