Table Of ContentProtozoa and Their Role in Marine Processes
NATO ASI Series
Advanced Science Institutes Series
A series presenting the results of activities sponsored by the NATO Science
Committee, which aims at the dissemination of advanced scientific and
technological knowledge, with a view to strengthening links between scientific
communities.
The Series is published by an international board of publishers in conjunction with
the NATO Scientific Affairs Division
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B Physics London and New York
C Mathematical and Kluwer Academic Publishers
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D Behavioural and
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E Applied Sciences
F Computer and Springer-Verlag
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G Ecological Sciences London Paris Tokyo Hong Kong
H Cell Biology Barcelona
Series G: Ecological Sciences Vol. 25
Protozoa and Their Role
in Marine Processes
Edited by
P.C. Reid
C. M. Turley
Plymouth Marine Laboratory
Citadel Hill
Plymouth, Devon PL 1 2PB
United Kingdom
and
PH. Burkill
Plymouth Marine Laboratory
Prospect Place
West Hoe
Plymouth, Devon PL 1 3DH
United Kingdom
Springer-Verlag Berlin Heidelberg New York
London Paris Tokyo Hong Kong Barcelona
Published in cooperation with NATO Scientific Affairs Division
Proceedings of the NATO Advanced Study Institute on Protozoa and Their Role in
Marine Processes held in Plymouth (United Kingdom), 24 July-5 August 1988.
ISBN-13: 978-3-642-73183-9 e-ISBN-13: 978-3-642-73181-5
DOl: 10.1007/978-3-642-73181-5
This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned,
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Copyright Law.
© Springer-Verlag Berlin Heidelberg 1991
Sof'tcover reprint of the hardcover 1s t edition 1991
2131/3140-543210 - Printed on acid-free-paper
PREFACE
In the summer of 1988, under NATO sponsorship, approximately 80
scientists lived and worked together in Plymouth for two weeks
to evaluate the ecological role of protozoa in the sea.
Through the convivial surroundings, close working conditions
and special facilities that had been brought together for NATO
ASI 604/87 a 'melting pot' of ideas was formed, which
stimulated the multidisciplinary creativity which is expressed
in this book and in a second series of papers which will be
published in Marine Microbial Food Webs under the title -
"Protozoa and their Role in Marine Microbial Food Webs".
Discussions of the role of protozoa in the microbial food web,
in the cycling of carbon and nitrogen and the extent to which
this web acts as a link or sink to metazoa in the water column
were major themes of the ASI. Structured sessions covering
oral and poster presentations, field work, model 1 ing,
laboratory practicals and demonstrations of techniques such as
image analysis and flow cytometry, formed the core of the
meeting. Participants took part enthusiastically in the
practical sessions developing new concepts and obtaining new
insights into their work. The practicals included a 'protozoo'
and some beautiful films and videos. Field excursions were
made to a range of sites including a unique marine sewage farm
at Looe in Cornwall, (Jones this volume). Interactive workshops
allowed scientists with no modelling experience to input their
results to three simulation models and a flow analysis package.
The major discussions of the ASI are encapsulated in the papers
of this book which comprise both invited keynote addresses and
session summaries. In the index the papers are grouped under
headings representing the main sessions of the meeting, which
in general, follow the order of presentation at the ASI.
The session on Taxonomy drew attention to the huge diversity of
'protists' found in marine environments, their varied
nutritional modes and outlined a revised evolutionary tree.
Few marine protists have so far been isolated in culture and
biotechnological applications are in their infancy, although
many of the results of the ASI have relevance in this area.
One presentation outlined the potential for manipulating the
microbial assemblage at the base of the food chain to maximise
yields in commercial prawn farming. Rapid advances are being
made in methodology, in particular, epifluorescence microscopy
has made a very great contribution to microbial studies since
the last ASI on a related subject in 1980 [Bougis P (ed) (1982)
Marine Pelagic Protozoa and Microzooplankton Ecology. Ann Inst
oceanogr Paris 58(S): 1-352]. This microscopic technique
contributed to one of the highlights of the meeting, the
recognition of the importance of chloroplast retention by
protists. Examples of endosymbiosis from a wide range of algal
groups were described, some retaining their original tests
within the host, others selectively adsorbing plastids. In the
sessions on Trophic Behaviour and Communi ty Grazing,
phagotrophic protists were seen to play an important role in
structuring food webs because of their fast growth rates and
VI
efficient energetics and production. Estimate of production
were seen to vary by two orders of magnitude. One study
reported selective grazing by heterotrophic nanoplankton on
bivalve gametes. This discovery has implications for
microzooplankton dynamics and may be one of the causes of year
to-year variability of intertidal settlement by gamete
releasing organisms. The session on Mineral Flux demonstrated
interannual variability of oceanic sedimentation from long-term
sediment trap studies - a discovery which has direct relevance
to studies of global climate change. The parallel session on
'Snow and Fluff', another highlight of the meeting, showed the
important role that protozoa play in the vertical transport of
particles in the sea and in the formation and decomposition of
aggregates in the water column and on arrival on the sea-bed.
Protozoa were also seen to act as 'crucial' agents in the
transfer and/or recycling of nutrients a theme which was
further developed in the modelling sessions. Ecological
discussions stressed the extreme diversity of habitats and
niches occupied by protozoa and their great temporal and
spatial variation. Larry Pomeroy, in his summation of the ASI,
using hierarchical theory, stressed the need to examine this
complexity at similar levels of ecological development.
The success of the meeting was only possible through the hard
work and enthusiasm of a dedicated team of local and
international helpers - see appended list of acknowledgements.
The local team included three sub-committees to cover
logistics, catering and the social programme. Delegation of
responsibilities to these committees, spread the workload,
broadened the input of ideas and proved essential to the
efficient running of the ASI. During the ASI Plymouth
celebrated the 400th anniversary of the Armada providing a
background and theme for a highly successful social programme
with Dave Robins as master of ceremonies.
I wish, in particular, to thank Peter Burkill for his
imagination and help in the planning and running of the ASI and
Carol Turley for her financial wizardry and great assistance in
the production of this book. The whole operation would not
have been possible without Angie Smith who ran the ASI
secretariat and typed, revised and concatenated this volume.
We are all indebted to Professor B L Bayne, Director, Plymouth
Marine Laboratory, for his support and for the use of
facilities during, prior and subsequent to the ASI. We also
thank Professor L Heath, Polytechnic South West, for access to
the well equipped laboratories of the Department of Biological
Sciences and for the enthusiastic support of his technicians.
The memories of this Plymouth ASI have left a lasting
impression on all participants which will, I believe, strongly
influence the future direction of marine microbial research.
It is our hope that this book will aid this process and provide
a fitting record of the hard work and team spirit engendered in
all who were involved.
Chris Reid
Director NATO ASI 604/87
ACKNOWLEDGEMENTS
The Director and Organising Committee of the NATO Advanced
Study Institute 604/87 on 'Protozoa and their Role in Marine
Processes' wish to acknowledge and thank NATO and the
following organisations, companies and individuals for their
support and assistance.
Commission of the European Communities
UNESCO
National Science Foundation, USA
Office of Naval Resarch, USA
Plymouth Marine Laboratory
Polytechnic South West
The Marine Biological Association of the UK
Plymouth City Council
Analytical Measuring Systems
The British Petroleum Company plc
Cell Systems
Coulter Electronics Ltd
Duncan and Associates
Ellis Horwood Ltd
Elsevier Applied Science Publishers Ltd
Nikon UK Ltd
Olympus Optical Co (UK) Ltd
St Austell Brewery Co Ltd
Sterilin Ltd
Vospers Rentals Ltd
Carl Zeiss (Oberkochen) Ltd
Main Committee Members: Local Organisers:
Dr PC Reid (Director) D Robins
Dr PH Burkill (Secretary) J Green
Dr CM Turley (Treasurer) A Smith
E Pilling
Other Committee Members: R Leakey
A John
Dr H Ducklow (USA) B Matthews
Dr JC Green (UK) P Radford
Dr MB Jones (UK) A Taylor
Prof F Rassoulzadegan (F) J Stephens
Dr Gene Small (USA) G Tarran
Prof V Smetacek (FRG) G Siley
M Wilson at Buckland Abbey
National Trust Staff at Cotehele
For preparation of manuscripts and their production in
camera-ready form, we thank Mrs Angie Smith, Dr Howard Bottrell
and Mrs Linda Jones, and for photographic assistance, Mr David
Nicholson.
CONTENTS
TAXONOMY
Taxonomy (46 - or more - protistan phyla) - Session summary
Eugene B Small and John C Green ........................... 1
A taxonomic review of heterotrophic protists important
in marine ecology
Michael A Sleigh .......................................... 9
METHODS
Methods for the study of marine microzooplankton -
Session summary
Fereidoun Rassoulzadegan 39
Quantitative sampling of field populations of
protozooplankton
Madhu A Paranj ape ......................................... 59
The application of image analysed fluorescence microscopy
for characterising planktonic bacteria and protists
Michael E Sieracki and Kenneth L Webb ..................... 77
Culturing marine protozoa - Session summary
Andrew J Cowling .......................................... 101
A method for the cloning and axenic cultivation of
marine protozoa
Anthony T Soldo and Sylvia A Brickson ..................... 105
POLLUTION
Pollution - Session summary
TorbjcPrn Dale ............................................. 113
Protists and pollution - with an emphasis on planktonic
ciliates and heavy metals
Torbj cPrn Dale ............................................. 115
Effects of saline sewage on the biological community of
a percolating filter
Malcolm B Jones and Ian Johnson ........................... 131
SYMBIOSIS
Endosymbiosis in the protozoa - Session summary
Michele Laval-Peuto ..........•............................ 143
Mixotrophy in marine planktonic ciliates: physiological
and ecological aspects of plastid-retention by
oligotrichs
Diane K Stoecker .....•.................•...•.............. 161
IX
AUTECOLOGY
Brief perspective on the autecology of marine protozoa
John J Lee ................................................ 181
TROPHIC BEHAVIOUR
Trophic behaviour - Session summary
Victor Smetacek ........................................... 195
COMMUNITY GRAZING
Community grazing in heterotrophic marine protista
Session summary
Gerard M Capriulo ......................................... 205
Trophic behaviour and related community feeding
activities of heterotrophic marine protists
Gerard M Carriulo, Evelyn B Sherr and Barry F Sherr ....... 219
ENERGETICS AND PRODUCTION
Protozoan energetics - Session summary
Johanna Laybourn-Parry .................................... 267
Global production of heterotrophic marine planktonic
ciliates
Dennis H Lynn and David JS Montagnes 281
MARINE 'SNOW' AND 'FLUFF'
Protozoa associated with marine 'snow' and 'fluff'
Session summary
Carol M Turley ............................................ 309
Protozoa as makers and breakers of marine aggregates.
Karin Lochte ...............................•....••.•...... 327
MINERAL FLUX
Mineral flux and biogeochemical cycles of marine
planktonic protozoa - Session summary
Kozo Takahashi ............................................ 347
Protista and mineral cycling in the sea
Barry SC Leadbea ter ....................................... 361
NUTRIENT CYCLING
Evolving role of protozoa in aquatic nutrient cycles
David A Caron ............................................. 387
Protozoans as agents in planktonic nutrient cycling
Tom Berman 417
x
MODELLING
Modelling - Session summary
Hugh W Ducklow and Arnold H Taylor ........................ 431
Modelling carbon and nitrogen flows in a microbial
plankton community
Colleen L Moloney and John G Field ........................ 443
WORKSHOP SUMMARY AND FORWARD LOOK
Status and future needs in protozoan ecology
Larry R Pomeroy ........................................... 475
LIST OF PARTICIPANTS ...................................... 493
SUBJECT INDEX ............................................. 498
