Table Of ContentEcological Studies, Vol. 127
Analysis and Synthesis
Edited by
M.M. Caldwell, Logan, USA
G. Heldmaier, Marburg, Germany
O.L. Lange, Wiirzburg, Germany
H.A. Mooney, Stanford, USA
E.-D. Schulze, Bayreuth, Germany
U. Sommer, Kiel, Germany
Ecologica! Studies
Volumes published since 1992 are listed at the end of this book.
Springer-V erlag Berlin Heidelberg GmbH
H. Sandermann A.R. Wellburn R.L. Heath (Eds.)
Forest Decline and Ozone
A Comparison of Controlled Chamber
and Field Experiments
With 77 Figures, 12 in Color and 22 Tables
t
Springer
Prof. Dr. Heinrich Sandermann
GSF-Forschungszentrum fur Umwelt und Gesundheit GmbH
Institut fur Biochemische Pfianzenpathologie
D-85764 OberschleiBheim, Germany
Prof. Dr. Alan R. Wellburn
University of Lancaster
Institute of Environmental and Biological Sciences
Lancaster LAI 4YQ, UK
Prof. Dr. Robert 1. Heath
U niversity of California, Riverside
Department of Botany and Plant Sciences
Riverside, CA 92521, USA
ISSN 0070-8356
ISBN 978-3-642-63912-8 ISBN 978-3-642-59233-1 (eBook)
DOI 10.1007/978-3-642-59233-1
Library of Congress Cataloging·in·Publication Data. Forest decline and ozone: a comparison of
controlled chamber and field experiments/Heinrich Sandermann, Alan R. Wellburn. Robert 1. Heath,
eds. p. cm. - (Ecologica! studies; voI. 127) Includes bibliographica! references and index. ISBN 3·
540·61321·8 1. Trees-Effect of atmospheric ozone on-United States. 2. Trees-Effect of atmospheric
ozone on-Europe. 3. Forest declines-United States. 4. Forest dec1ines-Europe. 1. Sandermann,
Heinrich. II. Wellburn, Alan, 1940· . III. Heath, Robert 1., 1940 . IV. Series: Ecological studies; v.
127. SB745.F67 1997 634.9'619-dc20 96·30571
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Preface
The idea for this book arose in 1993, after the Free State of Bavaria
through its Bayrisches Staatsministerium rur Landesentwicklung
und Umweltfragen (Bavarian Ministry of Regional Development
and the Environment) decided to discontinue both the Bavarian
project management (PBWU) for forest decline research and the
multidisciplinary field research on the Wank Mountain in the
Alps near Garmisch. Forest decline through the action of ozone
and other photooxidants was a main topic of the supported re
search in the Alps and will be a topic of new investigations in the
Bavarian Forest. Many interesting results were obtained, but the
researchers involved have not had sufficient time to allow reliable
conclusions to be drawn. It was therefore decided to ask inter
national experts for contributions in order to summarize the best
available evidence of a possible link between ozone and forest
decline - a topic which has been studied in the USA since the late
1950s and in Europe since the early 1980s.
The original idea of Waldsterben as an irreversible large-scale
dieback of forests in Germany was soon recognized to be wrong
(Forschungsbeirat 1989). However, the new criteria used for the
official German and European damage inventories (loss or yel
lowing of needles or leaves, tree morphology) indicate that per
sistently high percentages of damaged spruce and pine remain,
and there is an increasing percentage of damaged beech and oak,
with a high proportion of biotic disease (Forschungsbeirat 1989;
UN-ECE 1995). The enormous volume of damage inventory data
in Europe constitutes an operational definition of forest decline.
Yet, the symptoms evaluated are nonspecific and therefore diffi
cult to reproduce in laboratory experiments. Much better diag
nostics, including molecular stress marker techniques, are
needed. Two field guides to diagnose tree diseases have been
published: one for California where ozone problems were ori
ginally discovered (Scharpf 1993), and one for Germany in the
VI Preface
aftermath of the Waldsterben concept (Hartmann et al. 1988).
Both guides contain excellent color photographs and useful
glossaries with current definitions of decline, predisposition, and
other terms.
A functional definition of forest decline has been given by
Manion (1981). Decline was defined as a multiplicity of tree dis
eases due to more than a single cause. Manion (1981) also implied
that a strict temporal sequence of the abiotic and biotic agents
was involved, although this latter aspect may apply only in special
cases. The functional definition is close to the classical phyto
pathological concept of predisposition (Gaumann 1951; For
schungsbeirat 1989; Ayres 1984). In our case, abiotic stress factors
are thought to sensitize or desensitize a tree for biotic disease, and
vice versa. Forest decline phenomena have also been discussed on
the assumption of an abiotic origin (Huettl and Miiller-Dombois
1993). However, such a restricted definition does not appear to be
justified with respect to ozone since an interaction between ozone
exposure and fungal infection, as well as insect attack, has already
been established in the San Bernardino Mountains, California
(Scharpf 1993).
Atmospheric ozone is a relatively stable product of the pho
tooxidation of hydrocarbons and nitrogen oxides. It is phytotoxic
under two basic regimes. Firstly, acute toxicity occurs during
short-term exposure of plants to high ozone concentrations, and
symptoms are primarily caused by membrane disruption, cell
death, and tissue necrosis. Secondly, chronic toxicity occurs
during long-term exposure of plants to near-ambient ozone
concentrations. In this case, both direct and indirect symptoms
occur depending on stomatal apertures and interactions with
other environmental abiotic and biotic factors.
Conifers such as spruce (e.g., Picea abies) and pine (e.g., Pinus
ponderosa, Pinus sylvestris) are known to develop visible symp
toms primarily on the youngest and the older needle classes under
acute and chronic ozone exposure regimes, respectively (Hart
mann et al. 1988; Scharpf 1993). Due to moderate levels of ozone
in most forests, only the chronic exposure regime seems to be
ecologically relevant for conifers. The situation is less clear for
deciduous trees and crop plants. Prior to the development of
visible symptoms, however, biochemical stress reactions occur,
such as the development of ethylene or the induction of defence
reactions.
Preface VII
This book will attempt to describe {I} the temporal and causal
links between the initial stress reactions and visible ozone sym
ptoms; {2} the possible relationship between ozone effects and
complex tree diseases, and {3} the role of ozone in more general
forest ecosystem interactions.
At this time, definite proof of a link between ozone and forest
decline exists only for California. However, the new data sum
marized in this book starts to also support a role for ozone in
forest decline in eastern USA and Europe. Improved diagnostics
for the effects of ambient ozone still need to be developed in order
to prove or rule out a link between ozone and forest decline. The
present book therefore represents an interim report that illus
trates the complexities of ozone research and that hopefully will
stimulate more definitive research.
The idea for this volume is largely due to Prof. Dr. Dr. h.c.
Hubert Ziegler {Emeritus, Technische Universitat Miinchen} and
Prof. Dr. Ernst-Detlef Schulze {Universitat Bayreuth}. The con
stant interest and encouragement by the series editor of Ecolo
gical Studies, Prof. Dr. Dr. h.c. Otto 1. Lange {Emeritus,
Universitat Wiirzburg} are also gratefully acknowledged. Special
thanks are due to Drs. P. Miller, M. Arbaugh, and P. Temple who
organized an impressive excursion for the editors into the San
Bernardino Mountains to see first hand the longest-lasting study
site for ozone and the forest.
OberschleiBheim, September 1996 Heinrich Sandermann
References
Ayres PG (1984) The interaction between environmental stress injury and biotic
disease physiology. Annu Rev Phytopathol 22:53-75
Forschungsbeirat Waldschiiden/Luftverunreinigungen FBW (1989) Dritter Ber
icht, Kernforschungszentrum Karlsruhe GmbH, Karlsruhe, ISSN 0931-7805
Giiumann E (1951) Pflanzliche Infektionslehre. Birkhiiuser, Basel
Hartmann G, Nienhaus F, Butin N (1988) Farbatlas Waldschiiden. Diagnose von
Baumkrankheiten. E Wmer, Stuttgart
Huettl RF, Mueller-Dombois D (1993) Forest decline in the Atlantic and Pacific
regions. Springer Berlin Heidelberg New York, pp 351-358
VIII Preface
Manion PD (l981) Tree disease concepts. Prentice-Hall, Englewood Cliffs, pp
324-339
Scharpf RF (ed) (1993) Diseases of Pacific coast conifers. US Department of
Agriculture, Forest Service, Agricultural Handbook 521, Washington DC
UN-ECE (ed) (l995) Forest condition in Europe. United Nations Economic
Commission for Europe, Brussels, ISSN 1020-3729
Contents
1 Ozone Formation, Destruction and Exposure
in Europe and the United States
W.R. Stockwell, G. Kramm, H.-E. Scheel,
V.A. Mohnen and W. Seiler. . . . . . . . . . . . . . . . . 1
1.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 The Chemistry of Tropospheric Ozone Formation 4
1.3 Urban Ozone Formation and Transport
to Mountain Sites . . . . . . . . . . . . . . . . . . . . . . . . 10
1.4 Meteorological Effects
on Tropospheric Ozone Concentrations . . . . . . . . 11
1.5 Ozone Networks and the Analysis
of Measurements ........................ 14
1.6 The Global Distribution of Ozone Concentrations
and the Seasonal Cycle . . . . . . . . . . . . . . . . . . . . 17
1.7 Long-Term Trends
in Tropospheric Ozone Concentrations ........ 18
1.8 Present Ozone Concentrations
and Ozone Exposure over Europe
and North America. . . . . . . . . . . . . . . . . . . . . . . 24
1.9 Dry Deposition of Ozone .................. 27
1.10 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
References .. . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
2 Ozone and Its Known and Potential Effects
on Forests in Western United States
P.R. Miller, M.J. Arbaugh and P.J. Temple. . . . . . 39
2.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . .. 39
2.2 Spatial Distribution and Temporal Trends
of Ozone in Relation to Geographic Setting
and Summer Climate ..................... 40
x
Contents
2.2.1 Daily Patterns of Ozone Concentrations
in Rural Mountain Locations. . . . . . . . . . . . . . .. 42
2.2.2 Spatial Variation of Ozone in the Sierra Nevada
and San Bernardino Mountains . . . . . . . . . . . . .. 42
2.2.3 Temporal Trends of Ozone and the Outlook
for the Future. . . . . . . . . . . . . . . . . . . . . . . . . .. 42
2.3 History of Ozone Damage to Californian Forests 43
2.3.1 Field Identification of Ozone Injury. . . . . . . . . . . 43
2.3.2 Role of Other Biotic and Abiotic Stresses. . . . . . . 44
2.3.3 Crown Changes Associated
with Chronic Ozone Injury . . . . . . . . . . . . . . . . . 45
2.3.4 Spatial Distribution and Temporal Changes
of Ozone Damage . . . . . . . . . . . . . . . . . . . . . . .. 45
2.3.4.1 Southern California ...................... 45
2.3.4.2 Sierra Nevada .......................... 47
2.4 Experimental Exposures of Western Conifers
to Ozone. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
2.4.1 Foliar Injury Symptomatology. . . . . . . . . . . . . . . 48
2.4.2 Physiological Responses: Gas Exchange
and Photosynthesis. . . . . . . . . . . . . . . . . . . . . . . 50
2.4.3 Growth Responses of Seedlings and Saplings 52
2.5 Effects of Chronic Ozone Exposure
on Radial Growth in the Sierra Nevada
and San Bernardino Mountains . . . . . . . . . . . . .. 53
2.5.1 Seasonal Radial Growth Responses ........... 55
2.5.2 Long-Term Radial Growth Responses ......... 55
2.5.2.1 Direct Effects of Ozone on Radial Growth . . . . .. 55
2.5.2.2 Effects of Ozone Together with Climate
on Tree Growth . . . . . . . . . . . . . . . . . . . . . . . . . 57
2.5.3 Relationship Between Needle Retention
and Radial Growth Reductions .............. 57
2.5.4 Development and Composition
of Ozone-Exposed Forests
Under Euro-American Settlement Influences 58
2.5.4.1 Chronic Ozone Injury and Mortality
of Pines Due to Bark Beetle ................ 58
2.5.4.2 Ozone Influences on Recruitment
and Mortality of Ponderosa Pine Seedlings ..... 60
2.5.5 Status of Models for Simulating Ozone Effects
on Tree Growth and Stand Development . . . . . .. 60
2.6 Summary. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 61
References . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 63
