Arctic Ecosystems in a Changing Climate. An Ecophysiological Perspective

459 Pages·1991·12.03 MB·English

by  F. Stuart Chapin III

Physiological Ecology
A Series of Monographs, Texts, and Treatises
Series Editor
Harold A. Mooney
Stanford University, Stanford, California
Editorial Board
Fakhri Bazzaz F. Stuart Chapin James R. Ehleringer
Robert W. Pearcy Martyn M. Caldwell
KOZLOWSKI, Τ. T. Growth and Development of Trees, Volumes I and II,
1971
HILLEL, D. Soil and Water: Physical Principles and Processes, 1971
YOUNGER, V. B., and McKELL, C. M. (Eds.) The Biology and Utilization of
Grasses, 1972
KOZLOWSKI, Τ. T., and AHLGREN, C. E. (Eds.) Fire and Ecosystems, 1974
MUDD, J. B., and KOZLOWSKI, Τ. T. (Eds.) Responses of Plants to Air
Pollution, 1975
DAUBENMIRE, R. Plant Geography, 1978
LEVITT, J. Responses of Plants to Environmental Stresses, Second Edition.
Volume I: Chilling, Freezing, and High Temperature Stresses, 1980
Volume II: Water, Radiation, Salt, and Other Stresses, 1980
LARSEN, J. A. The Boreal Ecosystem, 1980
GAUTHREAUX, S. Α., JR. (Ed.) Animal Migration, Orientation, and
Navigation, 1981
VERNBERG, F. J., and VERNBERG, W. B. (Eds.) Functional Adaptations of
Marine Organisms, 1981
DURBIN, R. D. (Ed.) Toxins in Plant Disease, 1981
LYMAN, C. P., WILLIS, J. S., MALAN, Α., and WANG, L. C. H. Hibernation
and Torpor in Mammals and Birds, 1982
KOZLOWSKI, Τ. T. (Ed.) Flooding and Plant Growth, 1984
RICE, E. L. Allelopathy, Second Edition, 1984
List continues at the end of the volume.
Arctic Ecosystems in
a C h a n g i ng C l i m a te
An Ecophysiological Perspective
Edited by
F. S t u a rt C h a p in I II R o b e rt L. J e f f e r i es
Department of Integrative Biology Department of Botany
University of California, Berkeley University of Toronto
Berkeley, California Toronto, Ontario
Canada
J a m es F. R e y n o l ds G a i us R. S h a v er
Department of Botany Ecosystems Center
Duke University Marine Biological Laboratory
Durham, North Carolina Woods Hole, Massachusetts
J o s ef S v o b o da
Department of Botany
University of Toronto
Toronto, Ontario
Canada
E l l en W. C hu
Developmental Editor
Academic Press, Inc.
Harcourt Brace Jovanovich, Publishers
San Diego New York Boston London Sydney Tokyo Toronto
Front cover photo: Aerial view of a river drainage system with
hills of dwarf shrub tundra. For details see Chapter 4, p. 67 and
Color Plate 3.
This book is printed on acid-free paper. ©
Copyright © 1992 by ACADEMIC PRESS, INC.
All Rights Reserved.
No part of this publication may be reproduced or transmitted in any form or
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Academic Press, Inc.
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United Kingdom Edition published by
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Library of Congress Cataloging-in-Publication Data
Arctic ecosystems in a changing climate : an ecophysiological
perspective / F. Stuart Chapin III... [et al.].
p. cm. - (Physiological ecology series)
Includes index.
ISBN 0-12-168250-1
1. Plant physiological ecology-Arctic regions. 2. Vegetation and
climate-Arctic regions. 3. Vegetation dynamics-Arctic regions.
4. Global warming. I. Chapin, F. Stuart (Francis Stuart), iii.
II. Series.
QK474.A48 1991
581.5'2621-dc20 9120355
CIP
PRINTED IN THE UNITED STATES OF AMERICA
91 92 93 94 9 8 7 6 5 4 3 2 1
William Dwight Billings has provided inspiration, advice, exam­
ple, argument, and encouragement to generations of students of
ecology. He has played a leading role in the development of
physiological ecology over the past forty years, and his research
continues to lead the way to important advances in our under­
standing of the Arctic. Virtually every chapter in this book has
been influenced, directly or indirectly, by Dwight's contribu­
tions. It is thus with respect and affection that we dedicate this
book to William Dwight Billings.
Contributors
Numbers in parentheses indicate the pages on which the authors * contributions begin.
Frank Berendse (337), Centre for Agrobiological Research, NL-6700 AA
Wageningen, The Netherlands
W. D. Billings (91, 139), Department of Botany, Duke University, Durham,
North Carolina 29634
Caroline S. Bledsoe (301), Department of Land, Air, and Water Resources,
University of California, Davis, California 95616
L. C. Bliss (59, 111), Department of Botany, University of Washington, Seat-
tie, Washington 98195
John P. Bryant (377), Institute of Arctic Biology, University of Alaska, Fair­
banks, Alaska 99775
David M. Chapin (301), EA Engineering, Science, and Technology, Red­
mond, Washington 98052
F. Stuart Chapin ΠΙ (3, 321, 441), Department of Integrative Biology, Uni­
versity of California, Berkeley, Berkeley, California 04720
Todd E. Dawson (259), Section of Ecology and Systematics, Cornell Univer­
sity, Ithaca, New York 14853
Kaye R. Everett (35), Byrd Polar Research Center, Ohio State University,
Columbus, Ohio 43210
Ned Fetcher (359), Department of Biology, Faculty of Natural Sciences, Uni­
versity of Puerto Rico, Rio Piedras, Puerto Rico 00931
Τ. V. Gerasimenko (169), Laboratory of Ecology of Photosynthesis, Komarov
Botanical Institute, Academy of Sciences of the U.S.S.R., Leningrad 197022,
U.S.S.R.
Anne E. Giblin (281), The Ecosystems Center, Marine Biological Laboratory,
Woods Hole, Massachusetts 02543
S. Hahn (213), Lehrstuhl fur Botanik II, Universitat Wurzburg, D-8700
Wurzburg, Germany
G. Henry (391), Department of Geography, University of Alberta, Edmonton,
Alberta T6G 2E1, Canada
xiii
XIV Contnbutors
Larry D. Hinzman (35), Water Research Center, Institute of Northern Engi­
neering, University of Alaska, Fairbanks, Alaska 99775
Τ. I. Ivanova (169), Laboratory of Ecology of Photosynthesis, Komarov Botan­
ical Institute, Academy of Sciences of the U.S.S.R., Leningrad 197022,
U.S.S.R.
R. L. Jefferies (3, 391, 441), Department of Botany, University of Toronto,
Toronto, Ontario M5S 3B2
Svenjonasson (337), Institute of Plant Ecology, 0ster, Farimagsgade 2D, 1353
Copenhagen K, Denmark
Douglas L. Kane (35), Water Research Center, Institute of Northern Engi­
neering, University of Alaska, Fairbanks, Alaska 99775
Knut Kielland (321), Institute of Arctic Biology, University of Alaska, Fair­
banks, Alaska 99775
Jochen Kummerow (193), Department of Biology, San Diego State University,
San Diego, California 92182
O. L. Lange (213), Lehrstuhl fur Botanik II, Universitat Wurzburg, D-8700
Wurzburg, Germany
Paul W. Leadley (413), Department of Botany, Duke University, Durham,
North Carolina 27706
A. E. Linkins (281), Department of Biology, Clarkson University, Potsdam,
New York 13676
Ν. V. Matveyeva (59), Geobotanical Division, Komarov Botanical Institute, of
the Academy of Sciences of the U.S.S.R., Leningrad, 197022 U.S.S.R.
Barrie Maxwell (11), Canadian Climate Centre, Atmospheric Environment
Service, Downsview, Ontario M3H 5T4, Canada
James B. McGraw (359), Department of Biology, West Virginia University,
Morgantown, West Virginia 26506
Knute J. Nadelhoffer (281), The Ecosystems Center, Marine Biological Lab­
oratory, Woods Hole, Massachusetts 02543
Steven F. Oberbauer (213, 259), Department of Biological Sciences, Florida
International University, Miami, Florida 33199
Walter C. Oechel (139), Department of Biology, San Diego State University,
San Diego, California 92182
Κ. M. Peterson (111), Department of Biology, University of Alaska, Anchor­
age, Alaska 99508
M. Raillard (391), Department of Botany, University of Toronto, Toronto,
Ontario M5S 3B2, Canada
Paul B. Reichardt (377), Department of Chemistry, University of Alaska, Fair­
banks, Alaska 99775
J. F. Reynolds (3, 413, 441), Department of Botany, Duke University, Durham,
North Carolina 27706
Contributors XV
R. Ruess (391), Institute of Arctic Biology, University of Alaska, Fairbanks,
Alaska 99775
O. A. Semikhatova (169), Laboratory of Ecology of Photosynthesis, Komarov
Botanical Institute, Academy of Sciences of the U.S.S.R., Leningrad 197022,
U.S.S.R.
G. R. Shaver (3,193, 281, 441), Ecosystem Center, Marine Biology Laboratory,
Woods Hole, Massachusetts 02543
R. Siegwolf (213), Paul Scherrer Institut, Villigen (PSI), Switzerland
Bjartmar Sveinbjornsson (239), Department of Biological Sciences, University
of Alaska, Anchorage, Anchorage, Alaska 99508
J. Svoboda (3, 391, 441), Department of Botany, University of Toronto,
Toronto, Ontario M5S 3B2, Canada
J. D. Tenhunen (213), Systems Ecology Research Group, San Diego State Uni­
versity, San Diego, California 92182
Ming-ko Woo (35), Geography Department, McMaster University, Hamilton,
Ontario L8S 4L8, Canada
Preface
This book reviews the physiological ecology of arctic plants, suggests a new
role for physiological ecology in studying biotic controls over community and
ecosystem processes, and provides a physiological basis for predicting how
arctic plant communities will respond to global climate change. The Arctic
has always fascinated physiological ecologists because its rigorous environ­
ment challenges the physiological capabilities of organisms. For this reason,
physiological ecologists have, in the past, focused on the effects the physical
environment has on plants. In this book we review conclusions drawn from
this work but then go on to ask how physiological traits affect community and
ecosystem processes.
This approach is vital today, given the need to predict how the world's
ecosystems will respond to human-induced climatic change. Moreover, the
Arctic is expected to undergo the earliest and most pronounced effects of
global warming. In this book we examine the complex paths by which cli­
matic change might affect arctic plants and how these plant responses might
in turn influence arctic ecosystems and their potential feedbacks to the globe.
The book should thus prove useful to any practicing scientists, students of
ecology, and policy makers who are interested in the Arctic, physiological
ecology, ecosystem ecology, and global climate change.
The opening chapters describe the present and expected future environ­
ment of the Arctic, including climate, hydrology, and soils. The following
chapters discuss the diversity of arctic vegetation, its history, and the controls
over its limits at tree line. Chapters on individual physiological processes
examine how each process responds to the environment, how it influences
community and ecosystem processes, and how we might expect these con­
trols to change in response to altered climate. In the last chapter, the major
interactions and feedbacks between plants and their environment are con­
sidered as a whole in the light of a changing climate.
This book grew out of a conference funded by the National Science Foun­
dation and the United States Department of Energy. It is the first in a series that
will examine the responses of different ecosystems to global climate change.
xvii
1
Arctic Plant
Physiological Ecology:
A Challenge for the Future
F. S. Chapin III, R. L. Jefferies, J. F. Reynolds,
G. R. Shaver, and J. Svoboda
I. Introduction
II. Physiological Ecology and Ecosystem Studies
III. Physiological Ecology in the Arctic
IV. Climate Change: A Theme for Arctic Physiological
Ecology
References
I. Introduction
Many of the most exciting advances in knowledge occur when previously inde­
pendent disciplines merge. By integrating physiology and ecology, physio­
logical ecology has contributed substantially to our understanding of the
environmental controls over physiological processes and of the capacity of
organisms to tolerate different environments.
Although ecophysiological studies have been important from an evolu­
tionary perspective and offer insights into the mechanisms by which plants
and microbes cope with their environment, we suggest that the field has
reached a plateau and that most current research simply refines our un­
derstanding of various physiological processes. In the absence of a broader
context, physiological ecology can become a sterile series of "just-so" stories—
organisms live where they do because they have the physiological traits
enabling them to do so (Gould and Lewontin, 1979). Because the physiolog­
ical activities of plants and other organisms control many community and
ecosystem processes, community and ecosystem ecology can provide the
needed context for interpreting the importance of ecophysiological traits
beyond the level of the individual. Thus, studying the role of organisms in
Copyright © 1992 by Academic Press, Inc.
Arctic Ecosystems in a Changing Climate 3 All rights of reproduction in any form reserved.
4 Ε S. Chapin III et al.
controlling community and ecosystem processes should be a major research
direction in physiological ecology. Such an interdisciplinary approach to
ecosystem processes is imperative if we are to understand how impending
changes in climate will affect our globe.
II. Physiological Ecology and Ecosystem Studies
Physiological ecology has been used to predict ecosystem response to envi­
ronmental change in three primary ways, each of which has been criticized.
For physiological ecology to play a constructive role in ecosystem studies, the
limitations of the three approaches must be recognized and addressed:
1. Ecosystem response to environmental change is often predicted from
simple physiological responses of organisms (e.g., estimation of plant
production from temperature and light responses of photosynthesis). In
the long term, however, feedbacks among processes often govern perfor­
mance under natural circumstances more strongly than do short-term,
kinetic responses of individual processes. For example, even though pho­
tosynthesis always responds to C0 concentration in the short term, com­
2
pensatory changes in photosynthetic potential when plants are grown
under different C0 concentrations may counteract these short-term
2
effects (Tissue and Oechel, 1987; Chap. 7). Consequently, environmental
factors that exert strong short-term effects may not be influential over
longer time scales. To predict ecosystem response, we must study feed­
backs as well as direct environmental effects on plants.
2. Simulation modeling based on physiological studies can incorporate many
of the feedbacks that operate in natural ecosystems. Experience suggests,
however, that modeling predictions cannot be usefully extrapolated be­
yond two levels of organization (biochemistry, cell, leaf, canopy, popula­
tion, community, landscape, globe; Reynolds and Acock, 1985). For this
reason, the types of controls studied at one level of organization may not
be fully relevant at other levels. Most past ecophysiological work has
emphasized the direct effect of environment on physiology. Yet we need
to know the nature, strength, and timing of the feedbacks and time lags
that control resource supply and, indirectly, the growth of organisms. This
will require an integrated, whole-plant approach to physiological ecology.
Ecosystem and community ecology may provide the criteria for deciding
which feedbacks are important at these higher levels of organization.
3. Ecosystem response to environment has been predicted from comparisons
of current performance in two different environments (e.g., latitudinal
comparisons). A comparison of two ecosystems under equilibrium condi­
tions says nothing about the trajectory an ecosystem might follow in going
from one equilibrium state to another. Moreover, a new environment will
be occupied by different combinations of species than those co-occurring