Primary Productivity and Biogeochemical Cycles in the Sea

544 Pages·1992·25.586 MB·English

by  Richard W. Eppley (auth.), Paul G. Falkowski, Avril D. Woodhead, Katherine Vivirito (eds.)

Primary Productivity and
Biogeochemical Cycles in the Sea
ENVIRONMENTAL SCIENCE RESEARCH
Series Editor:
Herbert S. Rosenkranz
Department of Environmental and Occupational Health
Graduate School of Public Health
University of Pittsburgh
130 DeSoto Street
Pittsburgh, Pennsylvania
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Volume 43 - PRIMARY PRODUCTIVITY AND BIOGEOCHEMICAL CYCLES
IN THE SEA
Edited by Paul G. Falkowski and Avril D. Woodhead
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Primary Productivity and
Biogeochemical Cycles in the Sea
Edited by
Paul G. Falkowski
and
Avril D. Woodhead
Brookhaven National Laboratory
Upton, New York
Technical Editor:
Katherine Vivirito
Brookhaven National Laboratory
Upton, New York
SPRINGER SCIENCE+ BUSINESS MEDIA, LLC
Library of Congress Cataloging-in-Publication Data
Primary productivity and biogeochemica 1 cycles in the sea / edited by
Paul G. Falkowskl and AvMl D. Woodhead ; technical editor,
Katherine Vivirito.
p. cm. — (Environmental science research ; v. 43)
"Proceedings of the Conference on Primary Productivity and
Biogeochemica1 Cycles 1n the Sea, held June 2-6, 1991, at Brookhaven
National Laboratory, Upton, New York"—T.p. verso.
Includes bibliographical references and index.
ISBN 978-1-4899-0764-6
1. Marine productivity—Congresses. 2. Biogeochemical cycles-
-Congresses. I. Falkowskl, Paul G. II. Woodhead, Avril D.
III. Conference on Primary Productivity and Biogeochemica! Cycles in
the Sea (1991 : Brookhaven National Laboratory) IV. Series.
QH91.8.M34P75 1992
574.5'2636—dc20 92-3441
CIP
ISBN 978-1-4899-0764-6 ISBN 978-1-4899-0762-2 (eBook)
DOI 10.1007/978-1-4899-0762-2
Proceedings of the conference on Primary Productivity
and Biogeochemical Cycles in the Sea, held June 2-6, 1991,
at Brookhaven National Laboratory, Upton, New York
© Springer Science+Business Media New York 1992
Originally published by Plenum Press, New York in 1992
Softcover reprint of the hardcover 1st edition 1992
All rights reserved
No part of this book may be reproduced, stored in a retrieval system, or transmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming,
recording, or otherwise, without written permission from the Publisher
PREFACE
Biological processes in the oceans play a crucial role in regulating the fluxes of
many important elements such as carbon, nitrogen, sulfur, oxygen, phosphorus, and
silicon. As we come to the end of the 20th century, oceanographers have increasingly
focussed on how these elements are cycled within the ocean, the interdependencies of these
cycles, and the effect of the cycle on the composition of the earth's atmosphere and
climate. Many techniques and tools have been developed or adapted over the past decade
to help in this effort. These include satellite sensors of upper ocean phytoplankton
distributions, flow cytometry, molecular biological probes, sophisticated moored and
shipboard instrumentation, and vastly increased numerical modeling capabilities.
This volume is the result of the 37th Brookhaven Symposium in Biology, in which
a wide spectrum of oceanographers, chemists, biologists, and modelers discussed the
progress in understanding the role of primary producers in biogeochemical cycles. The
symposium is dedicated to Dr. Richard W. Eppley, an intellectual giant in biological
oceanography, who inspired a generation of scientists to delve into problems of
understanding biogeochemical cycles in the sea.
We gratefully acknowledge support from the U.S. Department of Energy, the
National Aeronautics and Space Administration, the National Science Foundation, the
National Oceanic and Atmospheric Administration, the Electric Power Research Institute,
and the Environmental Protection Agency. Special thanks to Claire Lamberti for her help
in producing this volume.
Symposium Committee
Paul Falkowski, Chairman
Robert Bidigare
Sally Chisholm
Dale Kiefer
Sharon Smith
v
CONTENTS
HONORARY LECTURE
Towards Understanding the Roles of Phytoplankton in Biogeochemical
Cycles: Personal Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . I
Richard W. Eppley
FACTORS LIMITING PRIMARY PRODUCTIVITY IN THE SEA
I. LIGHT
The Nature and Measurement of the Light Environment in the Ocean. . . . . 9
John T. O. Kirk
The Functional and Optical Absorption Cross-Sections of
Phytoplankton Photosynthesis. . . . . . . . . . . . . . . . . . . . . . . . . 31
Zvy Dubinsky
Molecular Ecology of Phytoplankton Photosynthesis . . . . . . . . . . . . . . . 47
Paul G. Falkowski
FACTORS LIMITING PRIMARY PRODUCTIVITY IN THE SEA
II. NUTRIENTS
Nutrient Limitation and Marine Photosynthesis . . . . . . . . . . . . . . . . . . 69
John Cullen, Xiaolong Yang, and Hugh L. MacIntyre
Geological and Climatic Time Scales of Nutrient Variability . . . . . . . . . . 89
Richard T. Barber
Nutrient Limitation of New Production in the Sea. . . . . . . . . . . . . . . . . 107
Richard Dugdale and Frances Wilkerson
Iron as a Limiting Factor in Oceanic Productivity. . . . . . . . . . . . . . . . . 123
John H. Martin
vii
ESTIMATION OF GLOBAL OCEAN PRODUCTION
Satellite Ocean Color Observations of Global Biogeochemical Cycles. . . . . 139
Marlon R. Lewis
Advances in Understanding Phytoplankton Fluorescence
and Photosynthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
Dale A. Kiefer and Rick A. Reynolds
Bio-Optical Models and the Problems of Scaling. . . . . . . . . . . . . . . . . . 175
Robert R. Bidigare, Barbara B. Prezelin, and Raymond C. Smith
TIIE ROLE OF MARINE ORGANISMS IN PRIMARY PRODUCTION
Phytoplankton Size. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Sallie W. Chisholm
Productivity of Seaweeds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
J. Ramus
Productivity of Zooxanthellae and Biogeochemical Cycles. . . . . . . . . . . . 257
Leonard Muscatine and Virginia Weis
NEW PRODUCTION AND BIOGEOCHEMICAL CYCLES
The Importance and Measurement of New Production. . . . . . . . . . . . . . 273
Trevor Platt, Pratima Jauhari, and Shubha Sathyendranath
The Role of Coastal High Latitude Ecosystems in Global
Export Production. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285
Paul K. Bienfang and David A. Ziemann
Tracer-Based Inferences of New Primary Production in the Sea . . . . . . . . 299
W.J. Jenkins and D.W.R. Wallace
New Production and the Global Carbon Cycle. . . . . . . . . . . . . . . . . . . 317
Jorge L. Sarmiento and Ulrich Siegenthaler
LOSS PROCESSES AND MATERIAL RECYCLING
Respiration: Taxation Without Representation? . . . . . . . . . . . . . . . . . . 333
Richard J. Geider
Bacterioplankton Roles in Cycling of Organic Matter:
The Microbial Food Web. . . . . . . . . . . . . . . . . . . . . . . . . . . 361
Jed Fuhrman
viii
Regeneration of Nutrients. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
William G. Harrison
Grazing, Temporal Changes of Phytoplankton Concentrations,
and the Microbial Loop in the Open Sea . . . . . . . . . . . . . . . . . . 409
Karl Banse
PHYTOPLANKTON IN TIIE GLOBAL CONTENT
Biosphere, Atmosphere, Ocean Interactions: A Plant
Physiologist's Perspective. . . . . . . . . . . . . . . . . . . . . . . . . . . 441
Joseph A. Berry
Reading the Sedimentary Record of the Ocean's Productivity. . . . . . . . . . 455
W.H. Berger and LC. Herguera
Do Marine Phytoplankton Influence Global Climate? . . . . . . . . . . . . . . 487
P.M. Holligan
Abstracts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505
Participants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 535
Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 545
ix
TOWARD UNDERSTANDING THE ROLES OF PHYTOPLANKTON
IN BIOGEOCHEMICAL CYCLES: PERSONAL NOTES
Richard W. Eppley
1969 Loring Street
San Diego, CA 92109
INTRODUCTION
This is a collection of remembrances of those who whetted my interest in
biogeochemical cycles in my formative years of the 1960s and 1970s. I used to ask myself
"How can anyone justify earning a living by studying phytoplankton?" Colleagues kept
pointing out that phytoplankton production for fish food was not a suitable justification.
The links between phytoplankton production and fish harvest, at least for local fisheries,
were not obvious. In search of more compelling reasons I began to lean on the role of
phytoplankton in C and N cycling as justification and to learn, especially from geochemists
and micropaleontologists, of the great importance of phytoplankton for all of us for all
time.
The people and events described are more or less in chronological order rather than
by subject matter. This order will lead to some awkwardness for the reader as will the
personal nature of this account. This is not intended to be a scientific report but rather
portions of a personal history. The pathway described starts in the fall of 1963 with my
arrival at the Scripps Institution of Oceanography (Scripps) and ends with the global
estimate of new production that Bruce Peterson and I reported in 1979 (Eppley and
Peterson, 1979).
EARLY DAYS IN THE FOOD CHAIN RESEARCH GROUP
Oceanography grew quickly in the early 1960s, resulting in the importation of many
people from elsewhere into the field; for example, I had studied seaweed physiology with
L. R. Blinks at Stanford's Hopkins Marine Station. The rapid growth of research funds
stopped in 1968 when Congress passed the Mansfield Amendment, an act that restricted
the funding of academic oceanography by the so-called "mission agencies" of the U. S.
government. Meanwhile, in 1962, Luigi Provasoli invited me to one of the AIBS
workshops on marine biology when the Soviet invitee was unable to attend. The workshop
included the leading phytoplankton researchers of that time: E. Steemann Nielsen, T.
Braarud, G. E. Hutchinson, G. A. Riley, G. E. Fogg, M. R. Droop, R. A. Vollenweider,
1. H. Ryther, and 10hn Strickland. As the junior researcher in the group, I listened a