Table Of ContentBrock/Springer Series in Contemporary Bioscience
Plankton
Ecology
Brock/Springer Series in Contemporary Bioscience
Series Editor: Thomas D. Brock
University of Wisconsin-Madison
Tom Fenchel
ECOLOGY OF PROTOZOA: The Biology of Free-living
Phagotrophic Protists
Johanna D6bereiner and Fabio o. Pedrosa
NITROGEN-FIXING BACTERIA IN NONLEGUMINOUS CROP
PLANTS
Tsutomu Hattori
THE VIABLE COUNT: Quantitative and Environmental Aspects
Roman Saliwanchik
PROTECTING BIOTECHNOLOGY INVENTIONS: A Guide for
Scientists
Hans G. Schlegel and Botho Bowien (Editors)
AUTOTROPHIC BACTERIA
Barbara Javor
HYPERSALINE ENVIRONMENTS: Microbiology and Biogeochemistry
Ulrich Sommer (Editor)
PLANKTON ECOLOGY: Succession in Plankton Communities
Ulrich Sommer (Ed.)
Plankton
Ecology
Succession in
Plankton Communities
With 104 Figures and 12 Tables
Springer-Verlag
Berlin Heidelberg New York
London Paris Tokyo
Library of Congress Cataloging-in-Publication Data
Plankton ecology: succession in plankton communtities / Ulrich Sommer
(ed.).
p. em. - (Brock/Springer series in contemporary bioscience)
Bibliography: p.
Includes index.
ISBN 978-3-642-74892-9
1: Plankton-Ecology. 2: Ecological succession. I. Sommer,
Ulnch, 1952- . II. Senes.
QH90.8.P5P53 1989
574'.92-dc20 89-11455
CIP
© 1989 by Springer-Verlag Berlin Heidelberg New York London Paris Tokyo.
Softcover reprint of the hardcover 1st edition 1989
All rights reserved. Neither this book nor any of its contents may be translated
or reproduced in any form, including electronic or digital means, without the
written permission of the publisher.
ISBN 978-3-642-74892-9 ISBN 978-3-642-74890-5 (eBook)
DOI 10.1007/978-3-642-74890-5
Production and editorial supervision: Science Tech Publishers
Cover art: Kandis Elliot
10 9 8 7 6 5 4 3 2 1
Contents
Preface vii
1 Toward a Darwinian Ecology of Plankton 1
Ulrich Sommer
2 Physical Determinants of Phytoplankton Succession 9
Colin S. Reynolds
3 The Role of Competition for Resources in 57
Phytoplankton Succession
Ulrich Sommer
4 The Role of Grazers in Phytoplankton Succession 107
Robert W. Sterner
5 The Role of Fungal Parasites in Phytoplankton 171
Succession
Ellen Van Donk
6 The Role of Competition in Zooplankton Succession 195
William R. DeMott
7 The Role of Predation in Zooplankton Succession 253
z.
Madej Gliwicz and Johanna Pijanowska
8 Toward an Autecology of Bacterioplankton 297
Carlos Pedros-Alio
9 The Role of Grazing on Bacteria in Plankton 337
Succession
Hans Giide
Index 365
v
Contributors
W.R. DeMott Department of Biological Sciences and Crooked Lake
Biological Station, Indiana University-Purdue University at Fort
Wayne, Fort Wayne, IN 46805, USA
Z.M. Gliwicz Department of Hydrobiology, University of Warsaw, Nowy
Swiat 67, 00-0046 Warsaw, Poland
H. Glide Institut fUr Seenforschung und Fischereiwesen, D-7994
Langenargen, FRG
C. Pedros-Alio Instituto de Ciencias del Mar, Consejo Superior
Investigaciones Cientificas, Paseo Nacional sin, 08003 Barcelona,
Spain
J. Pijanowska Department of Hydrobiology, University of Warsaw, Nowy
Swiat 67, 00-0046 Warsaw, Poland
C.S. Reynolds Freshwater Biological Association, Windermere
Laboratory, Ambleside, Cumbria, GB-LA22 aLP, UK
U. Sommer Max-Planck-Institute fUr Limnologie, P.O. Box 165, 2320
PIon, FRG
R.W. Sterner Department of Biology, UTA Box 19498, University of
Texas, Arlington, Arlington, TX 76109, USA
E. Van Donk Provincial Waterboard of Utrecht, P.O. Box 80300, 3508
TH Utrecht, The Netherlands
vii
Preface
hJ
"The lake as a microcosm", the title of one of the earliest papers limnology
(Forbes, 1887), could be the motto of this book. This book was written by
limnologists-especially plankton ecologists-for limnologists, marine biolo
gists, and general ecologists. It is centered around the mechanisms that drive
succession in plankton communities. There is no other field of limnology that
has been more influenced by population ecology. The concepts of population
ecology have been the skeleton to which plankton ecologists have attached
the empirical flesh. Now it is time to make the balance of trade even, and to
export the results of our work to the general ecological community. This book
is especially timely because many of the classic concepts of theoretical ecology
have recently come under criticism within the ecological community. We, the
plankton ecologists, have the empirical data, the experiments, and the case
studies that give life to the concepts of the theorists. Indeed, we conceive of
lakes, and especially their planktonic communities, as "microcosms", i.e., as
smaller models of the larger world of ecological interactions. This idea will
be further developed in Chapter 1 of this book.
Descriptions of individual successions and phytosociological classification
of succession patterns are of little interest for the nonplanktologist. However,
mechanisms that cause species replacements are of interest because plankton
do not differ fundamentally from other organisms in their requirements for
growth and survival. Only their need to remain in suspension is unique.
Otherwise, they face the same problems of fulfilling their nutritional demands
for growth and reproduction and of avoiding predation. As in any other
community, species-specific differences in the balance of reproduction and
mortality cause changes in species composition.
The requirements for nutrition, the consequences of being eaten, and the
functional division of the plankton community into phytoplankton, zooplank
ton, and bacterioplankton determine the order of this book. Chapters 2 to 5
deal with phytoplankton. In Chapter 2, the physical conditions (temperature,
turbulence, stratification, and light) necessary for their growth and survival
are explored; the stage is defined on which the successional play takes place.
Chapter 3 discusses the nutritional requirements of phytoplankton and the
successional consequences of competition for resources. The next two chapters
deal with the successional consequences of being food for others. Chapter 4
discusses the impact of zooplankton grazing, and Chapter 5 concerns the
impact of fungal parasitism. Chapters 6 and 7 cover the most important mech
anisms of zooplankton succession. Chapter 6 explores the role of food lim-
ix
x Preface
itation and competition for food, and Chapter 7 examines the impact of pre
dation by higher trophic levels.
The chapters on bacterioplankton are a bit different from the rest of the
book because the concept of succession as a series of species replacements
cannot-at least not yet-be applied to bacteria. Chapter 8 discusses the nu
tritional control of succession, metabolic guilds substituting for species. Growth
forms substitute for species in the treatment of predation on bacteria in Chap
ter 9.
The authors who have contributed to this book share the viewpoint that
succession in plankton can best be approached by understanding the adaptive
characters of the players and appreciating the importance of biotic interactions
that center around eating and being eaten. None of these views is universally
accepted and the debate continues. Hopefully, this book will provide an up
to-date and significant contribution to ongoing discussions of plankton ecology.
Acknowledgements: This book would not have been written without the
invitation by T.D. Brock to do so. This invitation was the stimulus that I
needed to summarize my own thoughts and work of the recent years and to
invite the other authors to do the same. I also want to express my gratitude
to all those with whom I have discussed plankton ecology during the recent
years, and who have helped me to develop my own view of the world of
plankton. In addition to those who are also contributors to this book, I want
to acknowledge especially S.S. Kilham, P. Kilham, W. Lampert, M. Tilzer, W.
Geller, and K.O. Rothaupt.
Ulrich Sommer
1
Toward a Darwinian Ecology
of Plankton
Ulrich Sommer
Max Planck Institute for Limnology
P16n, Federal Republic of Germany
The time now is ripe to summarize current views and hypotheses about the
mechanisms which drive succession in plankton communities, not only for
the sake of "understanding" plankton, but also in order to use the world of
plankton as a model for other kinds of ecological communities. Articles about
the seasonal changes in species composition of phyto-and zooplankton abound
in the literature. Unfortunately, they are mostly either descriptive or quite
speculative. Although the patterns of seasonality within individual bodies of
water can be quite regular, comparison between water bodies often leaves
the impression of chaos. Some authors draw a pessimistic conclusion and
consider the understanding of the mechanism of species shifts as an intractable
problem (Harris, 1986). If taken to the extreme, this point of view rejects the
idea that physiological ecology can be a legitimate enterprise in science. How
ever, other authors are still optimistic and continue to try the traditional
approach of the natural sciences to uncover causal relationships by the com
bined efforts of theoretical, experimental, and observational research. The
intellectual debate between the proponents of these two attitudes has become
increasingly hostile (d. Peters, 1986; Lehman, 1986) and has reached the
point where getting a referee from the opposing camp may seriously impair
the publication chances of a manuscript or the funding chances of a grant
proposal.
The authors of this volume, however, belong to the group of those who
have not given up. We still believe that it is possible to understand the mech
anisms driving species replacements. We derive this optimism from the sub
stantial progress in this area that has been made during the last decades
(Kerfoot, 1980; Reynolds, 1984; Lampert, 1985). This volume is intended to
summarize and review this progress. Therefore, we will not present a col-
1
2 Chapter 1 Toward a Darwinian Ecology of Plankton
lection of successional sequences or a typology of succession patterns for
different types of water bodies. Instead, we will suggest some mechanisms
that may drive species replacements in plankton communiities with the un
derstanding that these mechanisms operate in all kinds of pelagic environ
ments, although possibly with differences in relative importance. The reader
is invited to compare our reviews with earlier ones (e.g., in Hutchinson, 1967;
Smayda, 1980) to judge whether the progress since then justifies further hope
for clarifying the mechanism of succession in plankton communities.
It is the common opinion of the contributors to this book that we have
something to say not only to the plankton ecologists but also to ecologists in
general. Many of the divisive issues in plankton ecology are also debated in
general ecology (see Strong et al., 1983). The mechanisms discussed here are
traditional topics of physiological and population ecology (physical control,
nutrition, limitation of growth, competition, predation, parasitism). Many of
the classic issues of theoretical ecology can be brought into the focus of
experimental and field research more easily in plankton communities than
anywhere else. Representative samples of planktonic organisms can be easily
obtained. Estimates of abundance, biomass, and productivity are more reliable
than in any other community, despite the increasing awareness of plankton
patchiness. Cultivation of many plankton species is relatively easy and re
quires little space. Owing to their small body size, short generation times,
and high growth rates, space and time requirements for experiments with
them are modest, for example, in a I-liter container, a phytoplankton com
petition experiment can reach steady state in one to two months while several
years would be needed for herbs and several decades for trees. The small
body size of plankton organisms makes it possible to use experimental pop
ulations of millions of individuals, which excludes any unwanted influence
of inter-individual variability. The rapidity of population growth enables
successional sequences to be produced within one season that contain as many
stages as terrestrial vegetation succession does in the course of centuries. This
means that the plankton ecologist can make repeated direct observations
while the vegetation ecologist has to rely on indirect evidence from paleo
ecology or has to make the unlikely assumption that vegetation in different
places represents different stages of the same sequence.
Beginning in the early days of limnology, the seasonal cycles of plankton
abundances and species composition have been called "seasonal succession."
The usage of the term "succession" has been quite loose and the analogies
to the century-long, noncyclic process of terrestrial succession have rarely
been explored (however, see Reynolds, 1980). The word "succession" can
have two different meanings: the loose concept defines succession as a time
series of species replacements; the restrictive concept reserves the term succes
sion for those events in a chain of species replacements that are a consequence
of preceding ones (Reynolds, 1980). Species shifts in direct response to ex
ternal forcing factors do not qualify as succession. The restrictive definition
