Defense Mechanisms of Woody Plants Against Fungi

478 Pages·1992·12.93 MB·English

by  W. Merrill (auth.)

Springer Series in Wood Science
Robert A. Blanchette
Alan R. Biggs
Defense Mechanisms
of Woody Plants
Against Fungi

Springer Series in Wood Science
Editor: T. E. Timell
M. H. Zimmermann
Xylem Structure and the Ascent of Sap (1983)
J. F. Siau
Transport Processes in Wood (1984)
R.R. Archer
Growth Stresses and Strains in Trees (1986)
W.E. Hillis
Heartwood and Tree Exudates ( 1987)
S. Carlquist
Comparative Wood Anatomy (1988)
L. W. Roberts/P. B. Gahan/R. Aloni
Vascular Differentiation and Plant Growth Regulators (1988)
C. Skaar
Wood-Water Relations (1988)
J.M. Harris
Spiral Grain and Wave Phenomena in Wood Formation (1989)
B. J. Zobel/J.P. van Buijtenen
Wood Variation (1989)
P. Hakkila
Utilization of Residual Forest Biomass (1989)
J.W. Rowe
Natural Products of Woody Plants (1989)
K.-E. L. Eriksson I R. A. Blanchette I P. Ander
Microbial and Enzymatic Degradation of Wood and
Wood Components (1990)
R. A. Blanchette I A. R. Biggs
Defense Mechanisms of Woody Plants Against Fungi (1992)
S. Y. Lin/C. W. Dence
Methods in Lignin Chemistry (1992)
R. A. Blanchette A. R. Biggs (Eds.)
Defense Mechanisms
of Woody Plants
Against Fungi
With 209 Figures
Springer-Verlag Berlin Heidelberg GmbH
Professor Dr. ROBERT A. BLANCHETTE
Department of Plant Pathology
495 Borlaug Hall
1991 Upper Buford Circle
University of Minnesota
St. Paul, MN 55108, USA
Professor Dr. ALAN R. BIGGS
West Virginia University
Fruit Research Laboratory
P.O. Box 609
Kearneyville, WV 25430, USA
Series Editor:
Professor T. E. TIMELL
State University of New York
College of Environmental
Science and Forestry
Syracuse, NY 13210, USA
Cover: Transverse section of Pinus lambertiana wood. Courtesy of Dr. Cari de Zeeuw,
SUNY College of Environmental Science and Forestry, Syracuse, New York
ISBN 978-3-662-01644-2 ISBN 978-3-662-01642-8 (eBook)
DOI 10.1007/978-3-662-01642-8
Library of Congress Cataloging-in-Publication Data. Defense mechanisms of woody plants against
fungi / R. A. Blanchette, A. R. Biggs (eds.). p. cm. - (Springer series in wood science) Includes
bibliographical references and index. ISBN 3-540-54643-X (Springer-Verlag Berlin Heidelberg New
York). - ISBN 0-387-54643-X (Springer-Verlag New York Berlin Heidelberg) 1. Woody plants -
Disease and pest resistance. 2. 1fees - Disease and pest resistance. 3. Phytopathogenic fungi -
Host plants. 4. Fungal diseases of plants. /. Plant-fungi relationships. 1. Blanchette, R. A. (Robert
A.) II. Biggs, A.R. (Alan R.) III. Series. SB761.D35 1992 92-4617 582. 1' 5042326-dc20
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© Springer-Verlag Berlin Heidelberg 1992
Originally published by Springer-Verlag Berlin Heidelberg New York in 1992
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Preface
For the past decade, it has been apparent to both of us that a
reference text covering all aspects of tree defense mechanisms to fungi
was missing, needed and long overdue. Such a book would provide
a clear, comprehensive overview of how living roots, stems and leaves
respond to fungal pathogens. The need for such a book became in
creasingly clear to us from our conversations with each other, as well
as from our interactions with students and colleagues who desired a
sourcebook containing reviews of morphological, biochemical and
physiological aspects of host-parasite interactions in trees. During a
field trip sponsored by the Forest Pathology Committee of the Ameri
can Phytopathological Society, on a bus from one site to another, we
decided to take the responsibility to prepare a book of this type and
began to plan its composition.
To adequately address the topic of this book as we had envisioned
it, we believed that well-illustrated chapters were needed in order to
reflect the important advances made by the many investigators who
have examined the anatomical and physiological changes that occur
when trees are attacked by fungi. We are grateful to Dr. Tore Timell,
the Wood Science editor for Springer-Verlag, for supporting our
efforts and for providing an avenue to publish such a profusely il
lustrated volume. The Springer Series in Wood Science is one of the
finest quality book series published and readers of this book will find
a treasury of photomicrographs that document the changes that oc
cur in woody tissues in response to fungal pathogens. The numerous
illustrations and quality of the publishing are not without their price,
but the value they represent to those who use the book should surpass
the increased publication costs. We hope this book will serve as a
valuable reference to increase understanding of host-pathogen inter
actions and also to stimulate new interests that inspire continued
study in this exceedingly important area of tree pathology.
Within this book, readers will find chapters that address anatomi
cal and physiological responses of bark and xylem to mechanical in
juries and invasion by fungi. Response mechanisms in angiosperm
and gymnosperm roots, stems and leaves are presented in several
chapters that integrate information on how trees, as whole organisms,
defend and repair themselves. There are comprehensive chapters on
specific tree pathogens that have received considerable research atten
tion over the years, including Ophiostoma ulmi, Cronartium quer-
VI Preface
cuum f. sp. fusiforme and Hypoxylon mammatum. These diseases
have served, and continue to serve, as model systems for studying vas
cular wilts, rust and canker diseases. In addition, important factors
contributing to disease, such as plant-water relations and microen
vironmental aspects of wounded and colonized tissues, are discussed
within this volume. New approaches to studying tree defense reac
tions also are included and will be found in chapters covering molecu
lar, tissue culture and immunocytochemical aspects. How trees re
spond to fungi associated with insects and the role of mycorrhizae in
protecting tree roots against fungal pathogens are important con
tributions that expand the coverage of this book. The classic early
work has been included in the first chapter with a historical perspec
tive that reflects how much these early investigations still have to of
fer.
Researchers and students of tree pathology now h~ve a sourcebook
of information on defense mechanisms of woody plants against
fungi. Of prime importance, inquiring students have a reservoir of in
formation on how roots, stems and leaves respond to fungal patho
gens. We hope these contributions will be of great value and
assistance.
ROBERT A. BLANCHETTE and ALAN R. BIGGS
Contents
1 Mechanisms of Resistance to Fungi in Woody Plants:
A Historical Perspective
W. MERRILL
1.1 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.3 Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.4 Preformed Passive Barriers to Infection . . . . . . . . . . 4
1.4.1 Bark and Root Cortex . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.4.2 Cuticle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
1.4.3 Nature of the Cuticular Waxes . . . . . . . . . . . . . . . . . 5
1.4.4 Vessel Diameter and Spacing . . . . . . . . . . . . . . . . . . . 5
1.4.5 Heartwood Extractives . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.4.6 Xylem Decay Resistance . . . . . . . . . . . . . . . . . . . . . . . 6
1.4.7 Branch Stub Occlusion . . . . . . . . . . . . . . . . . . . . . . . . 6
1.4.8 Host Form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.5 Active Mechanisms of Resistance . . . . . . . . . . . . . . . 7
1.5.1 Walling-Out in the Cortex-Phloem . . . . . . . . . . . . . . 7
1.5.2 Walling-Out in the Xylem . . . . . . . . . . . . . . . . . . . . . . 7
1.5.3 Walling-Out in the Foliage . . . . . . . . . . . . . . . . . . . . . 8
1.5.4 Xylem "Double Banding" . . . . . . . . . . . . . . . . . . . . . . 8
1.5.5 Hypersensitive Responses . . . . . . . . . . . . . . . . . . . . . . 9
1.5.6 Second Party Interactions . . . . . . . . . . . . . . . . . . . . . . 9
1.6 An Iconoclastic Viewpoint . . . . . . . . . . . . . . . . . . . . . 10
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2 Anatomical and Physiological Responses
of Bark Tissues to Mechanical Injury
A.R. BIGGS
2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
2.2 Bark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.2.1 Tissues of the Inner Bark . . . . . . . . . . . . . . . . . . . . . . 14
2.2.2 Natural Periderms and Wound Periderms . . . . . . . . 16
2.3 Anatomy of Wound Response in Bark . . . . . . . . . . . 17
2.3.1 Light Microscopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.3.2 Electron Microscopy . . . . . . . . . . . . . . . . . . . . . . . . . . 23
2.4 Physiological and Biochemical Aspects of
Wounding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
VIII Contents
2.5 External Factors Influencing
the Generation of New Bark Tissues ........... . 30
2.6 Regeneration of Vascular Cambium
and Wound Closure .......................... . 32
2.7 Wounds as Infection Courts ................... . 34
2.8 Proposed Anatomical Model for Wound Responses
in Woody Plants ............................. . 35
References ........................................... . 36
3 Responses of Angiosperm Bark Tissues
to Fungi Causing Cankers and Canker Rots
A.R. BIGGS
3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
3.2 Infection Courts for Canker Pathogens . . . . . . . . . . 43
3.3 Colonization and Establishment
of Canker Pathogens . . . . . . . . . . . . . . . . . . . . . . . . . . 43
3.3.1 Valsa and Leucostoma Canker
of Peach and Various Hardwoods . . . . . . . . . . . . . . . 43
3.3.2 Nectria Canker of Apple and Various Hardwoods . 45
3.3.3 Hypoxylon Canker of Aspen . . . . . . . . . . . . . . . . . . . 45
3.3.4 Cryphonectria Canker of American Chestnut . . . . . 46
3.3.5 Eutypella Canker of Maple . . . . . . . . . . . . . . . . . . . . 46
3.3.6 Botryosphaeria Canker of Peach . . . . . . . . . . . . . . . . 47
3.3.7 Cerrena unicolor Canker Rot of Maple and Birch . 47
3.4 Delimitation of Canker Pathogens in Bark Tissues 47
3.5 Breaching of Host Boundaries
by Canker Pathogens . . . . . . . . . . . . . . . . . . . . . . . . . . 52
3.6 Expression of Resistance to Canker Pathogens
in Bark Tissues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
3.7 Molecular Regulation of Plant Response:
Normal Development, Wounding,
and Pathogenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
4 Responses of Gymnosperm Bark Tissues
to Fungal Infections
S. WOODWARD
4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
4.2 Physical Barriers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
4.2.1 Structure of the Periderm and Bark Cortex . . . . . . 62
4.2.2 Role of Suberized and Lignified Tissues
in Preventing Entry of Potential Pathogens . . . . . . . 63
4.3 Chemical Barriers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
4.3.1 Antifungal Phenolic Compounds in the Bark . . . . . 64
4.3.2 Resin Terpenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Contents IX
4.4 Responses of Bark to Wounding
and Pathogen Attack ......................... . 67
4.4.1 Physical Alterations, "Walling-Off" of Wounds
and Restoration of the Periderm
(Necrophylactic Periderm Formation) ........... . 67
4.4.1.1 Responses to Necrotrophic Pathogens .......... . 67
4.4.1.2 Responses to Biotrophic Pathogens ............. . 70
4.4.2 Role of Antimicrobial Chemicals
in the Wound Healing Response ............... . 70
4.5 Conclusions ........... 71
0 0. 0 •••••• 0 ••• 0. 0 ••••••
References .... 72
0 ••• 0 0 ••••••••• 0 • 0 • 0 • 0 ••••••••••••••••••
5 Anatomical Responses of Xylem to Injury and Invasion
by Fungi
R. A. BLANCHETTE
5.1 Introduction ......... 0. ......... 0............ 76
5.2 Anatomy of Sound Wood .. 0. ......... 0........ 76
5.3 Changes in Preexisting Xylem After Wounding . . . 77
5.4 Compartmentalization . 0. ... 0.................. 81
5.5 Formation of the Barrier Zone . . . . . . . . . . . . . . . . . 86
5.6 Xylem Decay Processes . . . . . . . . . . . . . . . . . . . . . . . . 89
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
6 Microenvironmental Aspects of Xylem Defenses
to Wood Decay Fungi
L. BODDY
6.1 Introduction 96
6.2 Xylem Microenvironment ..................... . 96
6.2.1 Resource Composition ........................ . 96
6.2.1.1 Xylem Anatomy: Routes of Access ............. . 96
6.2.1.2 Nutrients ... 98
0 ••••••••••••••••••••••••••••••••
6.2.1.3 Extractives ................. 98
0 •••••••••••• 0 0 • 0 •
6.2.1.4 Wood Acidity .. 99
0 •••••••• 0 ••••••••••••• 0 •••• 0 •
6.2.2 Microclimate: Water and Gaseous Regime .. 99
0 •••••
6.2.2.1 Problems of Measurement and Expression
of Water Content ............................ . 99
6.2.2.2 Gaseous Regime and Distribution of Water
in Xylem ................................... . 100
6.2.3 Effect of Wounding on Microclimate ........... . 102
6.2.4 Effect of Microbial Colonization on Microclimate 104
6.2.4.1 Heartwood ............................. 105
0 0 ••••
6.2.4.2 Sapwood . 105
0 ••••••• 0 ••• 0 0 •• 0. 0 •• 0 •••••••••• 0 •••
6.3 Effect of Microenvironment on Development
of Fungi in Xylem .... 106
0 0 ••••••••••••••••••••••