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Lipids in Aquatic Ecosystems
Michael T. Arts • Michael T. Brett
Martin J. Kainz
Editors
Lipids in Aquatic Ecosystems
ISBN: 978-0-387-88607-7 e-ISBN: 978-0-387-89366-2
DOI: 10.1007/978-0-387-89366-2
Springer Dordrecht Heidelberg London New York
Library of Congress Control Number: 2008942065
© Springer Science+Business Media, LLC 2009
All rights reserved. This work may not be translated or copied in whole or in part without the written
permission of the publisher (Springer Science+Business Media, LLC, 233 Spring Street, New York,
NY 10013, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in
connection with any form of information storage and retrieval, electronic adaptation, computer software,
or by similar or dissimilar methodology now known or hereafter developed is forbidden.
The use in this publication of trade names, trademarks, service marks, and similar terms, even if they
are not identified as such, is not to be taken as an expression of opinion as to whether or not they are
subject to proprietary rights.
The artwork depicted in the small inset on the front cover is a collaboration between the three editors
and the artist, Andrew Turnbull (www.turnbullsculpture.com), with subsequent modifications by graphic
artist Lucas Neilson.
Printed on acid-free paper
Springer is part of Springer Science+Business Media (www.springer.com)
Editors
Michael T. Arts
Aquatic Ecosystems Management
Research Division
National Water Research Institute –
Environment Canada
P.O. Box 5050, 867 Lakeshore Road
Burlington, ON, Canada L7R 4A6
Michael T. Brett
Department of Civil & Environmental
Engineering
University of Washington
Box 352700, 301 More Hall, Seattle
WA 98195-2700, USA
Martin J. Kainz
WasserKluster Lunz
Biologische Station
Dr. Carl Kupelwieser Promenade 5
3293 Lunz am See, Austria
Foreword
The direction of science is often driven by methodological progress, and the topic
of this book is no exception. I remember sitting with a visitor on the terrace of a
hotel overlooking Lake Constance in the early 1970s. We were discussing the gravimetric method of measuring total lipids in zooplankton. A few years later, as a visitor in Clyde E. Goulden’s lab, I was greatly impressed by the ability of an
instrument called an Iatroscan to discriminate and quantify specific lipid classes
(e.g., triacylglycerols, polar lipids, wax esters). At that time, food web analysis was
mainly concerned with bulk quantitative aspects. For example, lipids, because of
their high energy content, were considered mainly as an important food source and
storage product.
Nearly a decade ago, when Michael Arts and Bruce Wainman edited the first
volume entitled “Lipids in Freshwater Ecosystems” (Springer), the focus had
already changed. Fatty acid analysis had become more mainstream, because
new, less expensive, instruments had become available for ecological laboratories and because ecology, in general, was diversifying and integrating with other
disciplines. Hence, there was increased emphasis on studies which dealt with the
qualitative aspects of lipid composition. The concept of lipids in ecosystems was
no longer restricted to just providing fuel; lipid composition had, by then,
already been recognized as a factor controlling the flow of matter and the structure of food webs. In his foreword to the first book, Robert G. Wetzel defined a
rapidly evolving field that he called “biochemical limnology” and identified
lipid research as one of its facets. Judging from the ever increasing numbers of
published papers and congress contributions the field is presently evolving even
more rapidly.
However, progress was not restricted to limnology. In fact, methods of lipid and
fatty acid analysis were probably more advanced in marine ecology, and essential
fatty acids were an important factor in marine aquaculture. Lipid research in aquatic
organisms profited also from the growing connections to human nutrition science
interested in the importance of highly unsaturated fatty acids (HUFA; fatty acids
with ³20 carbons and ³3 double bonds) originating from fish and shellfish.
This became very evident at the 2002 summer meeting of the American Society of
Limnology and Oceanography in Victoria, British Columbia, when Michael
A. Crawford delivered an unusual, but fascinating plenary lecture entitled
v
“The evolution of the human brain.” Consequently, this new volume has broadened
its scope from freshwater to “aquatic ecosystems.” It is, thus, a contribution to finding common principles in marine and freshwater systems.
My personal interest in fatty acids has been stimulated again in recent years by
the controversy over food quality factors controlling the growth of zooplankton,
which used to be more a topic in limnology than in marine ecology. Two schools
developed at about the same time, one proposing that zooplankton growth was
limited by the availability of essential fatty acids, the other one developing the
concept of zooplankton growth controlled by inorganic nutrient stoichiometry. In
principle, both groups of resources can be limiting as they must be taken up with
the same food package and cannot be completely synthesized by the consumer
itself. Unfortunately, the empirical data were contradictory, and there was support
for both concepts. As usual, this resulted in a heated debate; however, we are now
on the way to a concept incorporating both groups of resources as limiting factors.
The controversy had a striking effect on aquatic lipid research; it stimulated discussion, created new ideas, and fostered methodological progress. Lipids and fatty
acids are now regular topics of special sessions at aquatic science conferences.
Robert Wetzel’s statements in the earlier foreword are still valid and up-to-date,
but the field has broadened considerably in the past decade. The “classical” studies
on lipids as storage products and carriers of lipophilic contaminants are continuing.
Research on lipids as nutritional factors now concentrates on the role of essential
components, e.g., polyunsaturated fatty acids (PUFA) and sterols, in modifying the
growth and reproduction of animals. This includes studies on biosynthesis and
metabolic pathways in food organisms and the characterization of fatty acid profiles
in organisms at the base of food webs and in allochthonous material. Spatial and
temporal variations in lipid composition need to be investigated to reach the goal of
a mechanistic prediction of food web structures under changing environmental
conditions. Finally, specific fatty acids and ratios of fatty acids are being developed
as biomarkers to aid in the identification of key food web connections.
Evolutionary ecology is beginning to explore adaptations of organisms to the
changing availability of essential fatty acids in their food, e.g., the evolution of life
histories, provision of offspring with PUFA, and the timing of diapause. However,
lipid production may also be considered as an adaptation by algae and bacteria
against their consumers. Evidence is accumulating indicating that not all fatty acids
are beneficial to consumers. Some are toxic or are precursors of toxic products, and
the question therefore now arises as to why organisms produce such costly
products.
Finally, lipid and fatty acid research has gained considerable applied importance
as humans are often “top predators” and also depend on essential dietary nutrients.
Public awareness of healthy nutrition is increasing, and this relates to both acquiring necessary food compounds and avoiding toxic contaminants. Lipids play a key
role in these processes.
The past 10 years have seen a rapid increase in our knowledge about the ecological importance of lipids. As with all progressive scientific initiatives this new
knowledge has also generated new questions. It is thus time for a new synthesis.
vi Foreword
This book addresses most of the topics mentioned above; hence it is a timely
book. I am sure it will not only summarize the status quo; it will also stimulate
new research within the important and exciting field of biochemical aquatic ecology as well as foster new and fruitful connections with the field of human
nutrition.
Plön, Germany Winfried Lampert
Foreword vii
Introduction .................................................................................................... xv
Michael T. Arts, Michael T. Brett, and Martin J. Kainz
1 Algal Lipids and Effect of the Environment
on their Biochemistry .............................................................................. 1
Irina A. Guschina and John L. Harwood
2 Formation and Transfer of Fatty Acids in Aquatic
Microbial Food Webs: Role of Heterotrophic Protists ......................... 25
Christian Desvilettes and Alexandre Bec
3 Ecological Significance of Sterols in Aquatic Food Webs .................... 43
Dominik Martin-Creuzburg and Eric von Elert
4 Fatty Acids and Oxylipins as Semiochemicals ...................................... 65
Susan B. Watson, Gary Caldwell, and Georg Pohnert
5 Integrating Lipids and Contaminants in
Aquatic Ecology and Ecotoxicology ....................................................... 93
Martin J. Kainz and Aaron T. Fisk
6 Crustacean Zooplankton Fatty Acid Composition ............................... 115
Michael T. Brett, Dörthe C. Müller-Navarra, and Jonas Persson
7 Fatty Acid Ratios in Freshwater Fish, Zooplankton
and Zoobenthos – Are There Specific Optima? .................................... 147
Gunnel Ahlgren, Tobias Vrede, and Willem Goedkoop
8 Preliminary Estimates of the Export of Omega-3
Highly Unsaturated Fatty Acids (EPA + DHA) from
Aquatic to Terrestrial Ecosystems.......................................................... 179
Michail I. Gladyshev, Michael T. Arts, and Nadezhda, N. Sushchik
Contents
ix
9 Biosynthesis of Polyunsaturated Fatty Acids in Aquatic
Ecosystems: General Pathways and New Directions ......................... 211
Michael V. Bell and Douglas R. Tocher
10 Health and Condition in Fish: The Influence of Lipids
on Membrane Competency and Immune Response ........................... 237
Michael T. Arts and Christopher C. Kohler
11 Lipids in Marine Copepods: Latitudinal Characteristics
and Perspective to Global Warming .................................................... 257
Gerhard Kattner and Wilhelm Hagen
12 Tracing Aquatic Food Webs Using Fatty Acids:
From Qualitative Indicators to Quantitative Determination ............ 281
Sara J. Iverson
13 Essential Fatty Acids in Aquatic Food Webs ...................................... 309
Christopher C. Parrish
14 Human Life: Caught in the Food Web ................................................ 327
William E. M. Lands
Name Index ..................................................................................................... 355
Subject Index .................................................................................................. 367
x Contents
Contributors
Gunnel Ahlgren
Department of Ecology and Evolution (Limnology), Uppsala University,
P.O. Box 573, 751 23 Uppsala, Sweden
Michael T. Arts
Aquatic Ecosystems Management Research Division, National Water Research
Institute – Environment Canada, P.O. Box 5050, 867 Lakeshore Road,
Burlington, ON, Canada L7R 4A6
Alexandre Bec
Laboratoire de Biologie des Protistes, Université Blaise Pascal,
Clermont-Ferrand II, Campus des Cézeaux, 63177 Aubiere Cedex, France
Michael V. Bell
Institute of Aquaculture, University of Stirling, Stirling, Stirlingshire FK9 4LA, UK
Michael T. Brett
Department of Civil & Environmental Engineering, University of Washington,
Box 352700, 301 More Hall, Seattle, WA 98195-2700, USA
Gary Caldwell
School of Marine Science and Technology, Newcastle University, Ridley
Building, Rm 354, Claremont Road, Newcastle upon Tyne NE1 7RU, UK
Christian Desvilettes
Laboratoire de Biologie des Protistes, Université Blaise Pascal,
Clermont-Ferrand II, Campus des Cézeaux, 63177 Aubiere Cedex, France
xi
Aaron T. Fisk
Department of Biology (Great Lakes Institute for Environmental Research),
University of Windsor, 2990 Riverside Drive West, Windsor, ON,
Canada N9B 2P3
Michail Gladyshev
Institute of Biophysics, Siberian Branch of the Russian Academy of Sciences,
660036 Krasnoyarsk, Akademgorodok, Russia
Willem Goedkoop
Department of Environmental Assessment, Swedish University of Agricultural
Sciences, Box 7050, 750 07 Uppsala, Sweden
Martin Graeve
Pelagic Ecosystems/Marine Chemistry and Marine Natural Products,
Alfred Wegener Institut für Polar- und Meeresforschung, Am Handelshafen 12,
27570 Bremerhaven, Germany
Irina A. Guschina
School of Biosciences, Cardiff University, P.O. Box 911, Cardiff CF10 3US,
Wales, UK
Wilhelm Hagen
Marine Zoology (FB2), Universität Bremen, P.O. Box 330440,
28334 Bremen, Germany
John L. Harwood
School of Biosciences, Cardiff University, P.O. Box 911, Cardiff CF10 3US,
Wales, UK
Sara Iverson
Department of Biology – Life Sciences Centre, Dalhousie University,
1355 Oxford Street, Halifax, NS, Canada B3H 4J1
Martin J. Kainz
WasserKluster Lunz – Biologische Station, Dr. Carl Kupelwieser Promenade 5,
A-3293 Lunz am See, Austria
xii Contributors
Gerhard Kattner
Pelagic Ecosystems/Marine Chemistry and Marine Natural Products,
Alfred Wegener Institut für Polar- und Meeresforschung, Am Handelshafen 12,
27570 Bremerhaven, Germany
Christopher C. Kohler
Director, Fisheries and Illinois Aquaculture Center, Southern Illinois University,
Carbondale, IL 62901-6511 USA,
Winfried Lampert
Max Planck Institute for Limnology, Plön, Germany
William E. M. Lands
6100 Westchester Park Drive, Apt. #1219, College Park, MD 20740, USA
Dominik Martin-Creuzburg
Limnological Institute, Universität Konstanz, Mainaustrasse 252,
78464 Konstanz, Germany
Dörthe Müller-Navarra
Aquatic Ecology, Universität Hamburg, Zeiseweg 9, 22609 Hamburg, Germany
Christopher C. Parrish
Ocean Sciences Centre, Memorial University of Newfoundland,
St. John’s, NF, Canada A1C 5S7
Jonas Persson
Department of Ecology and Evolution, Uppsala University, Husargatan 3,
75 123 Uppsala, Sweden
Georg Pohnert
Laboratory of Chemical Ecology – LECH, Ecole Polytechnique Fédérale
de Lausanne, EPFL SB ISIC LECH – BCH 4306, 1015 Lausanne, Switzerland
Nadezhda N. Sushchik
Institute of Biophysics, Siberian Branch of the Russian Academy of Sciences,
660036 Krasnoyarsk, Akademgorodok Russia,
Contributors xiii
Douglas R. Tocher
Institute of Aquaculture, University of Stirling, Stirling, Stirlingshire FK9 4LA, UK
Eric von Elert
Institute of Zoology, Universität zu Koeln, Weyertal 119, 50923 Koeln, Germany
Tobias Vrede
Department of Ecology and Environmental Sciences, Umeå University,
90187 Umeå, Sweden
Susan B. Watson
Aquatic Ecosystems Management Research Division, National Water Research
Institute – Environment Canada, P.O. Box 5050, 867 Lakeshore Road,
Burlington, ON, Canada L7R 4A6
xiv Contributors
Lipids in Aquatic Ecosystems
Michael T. Arts, Michael T. Brett , and Martin J. Kainz
Introduction
Life began as a process of self-organization within a lifeless environment. For single and, subsequently, multicellular organisms to differentiate themselves from the
outside world, they needed an effective, adaptable barrier (i.e., the cell/cytoplasmic
membrane). The modern cell membrane is mainly composed of phospholipids,
proteins, and sterols, which in unison regulate what goes into and out of the cell.
Some have hypothesized that spontaneously formed phospholipid bilayers played a
key role in the origin of life. The precise structure and composition of these biochemical groups have an enormous influence on the integrity and physiological
competency of the cell. It should not be surprising that this organizational and
functional specificity at the cellular level readily translates into profound systemic
effects at the macroscopic level. Thus, cellular lipid composition and organization
orchestrate both subtle and obvious effects on the health and function of organisms
→ populations → communities → ecosystems.
Ecology is, by its very nature, an integrative field of inquiry that actively promotes the examination of processes that span both cellular and macroscopic levels
of organization. Modern ecologists are challenged and motivated to put their
research into a broader perspective; ecology thrives at the intersections of disciplines! Lipids provide an effective platform for this mandate because they are a
global energy currency and because of their far-reaching physiological roles in
aquatic and terrestrial biota. Two previous, comprehensive efforts to examine the
role of lipids in aquatic environments exist. The first (Gulati and DeMott 1997)
arose as the proceedings of an international workshop held at Nieuwersluis, the
Netherlands in 1996. The objective of this workshop was “to take stock of the state
of the art in food quality research, to address factors that determine food quality”
and “to integrate the available information into a coherent and consistent view of
xv
M.T. Arts (), M.T. Brett , and M.J. Kainz
Aquatic Ecosystems Management Research Division , National Water Research Institute –
Environment Canada , P.O. Box 5050, 867 Lakeshore Road , Burlington , ON , Canada L7R 4A6
e-mail: [email protected]
food quality for the zooplankton.” A second, more extensive publication followed
2 years later (Arts and Wainman 1999) . That publication set about to “establish a
general reference and review book for those interested in aquatic lipids” and to
“demystify lipid research.” Its focus was mainly on freshwater ecosystems. Since
these two publications in the late 1990s, the field has advanced considerably, most
notably in such areas as:
• Refining the understanding of the essentiality of specific lipids
• Biochemical pathways and controls on PUFA synthesis and degradation
• Fatty acid as trophic markers
• Importance/essentiality of sterols
• Integrating contaminant and lipid pathways
• Trophic upgrading by protists, heterotrophic flagellates, and zooplankton
• Role of fatty acids and other lipids in the maintenance of membrane fluidity
• Role of fatty acids in cell signaling
• Effect of essential fatty acids (EFAs) on human health and behavior (e.g., n-3
deficiency)
• EFAs as seen from a conservation perspective
Advances such as these convinced us that, nearly a decade after the first edition, a
second book project should be undertaken. We envisioned that this book should (a)
have a much broader mandate than the original; for example, it should encompass
both freshwater and marine ecosystems, (b) touch on several of the recent advances
highlighted above, and (c) break new ground by interconnecting the fields of lipid
research with other highly topical areas such as climate change, conservation, and
human health.
A survey of the literature clearly shows that interest in lipids within environmental sciences is increasing almost exponentially. As more detailed and informative
experiments and observations are made, it is becoming clear that some lipids (e.g.,
the long chain, polyunsaturated, omega-3 fatty acid “docosahexaenoic acid” or
“DHA” for short, 22:6n-3) have a critical role to play in maintaining the health and
functional integrity of both aquatic and terrestrial organisms. Thus, the more general
interest in lipids as structural components and as purveyors of energy is increasingly
being coupled with this deeper understanding resulting in a parallel increase in publications dealing specifically with individual lipid molecules such as DHA.
The chapters in this book are broadly organized so as to elaborate and synthesize
concepts related to the role of lipids from lower to higher trophic levels up to and
including humans – an objective that has seldom been attempted from an ecological
perspective. A précis of the book’s 14 chapters follows:
In Chap. 1, “Algal Lipids and Effect of the Environment on Their Biochemistry,”
Irina Guschina and John Harwood explore the origins and synthesis of a wide variety of algal lipids (glycolipids, phospholipids, betaine lipids, and nonpolar glycerolipids) and provide important clues as to how environmental signals (temperature,
light, salinity, and pH) may influence the production of specific lipids and lipid
classes. Their chapter concludes with a concise summary of how nutrients and
nutrient regimes affect the production of lipids in algae.
xvi Introduction