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.