The Vascular Cambium: Development and Structure

736 Pages·1994·24.389 MB·English

by  Dr. Philip R. Larson (auth.)

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)
e.
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 (Ed.)
Natural Products of Woody Plants (1989)
K.-E.L. Eriksson/R. A. Blanchette/P. Ander
Microbial and Enzymatic Degradation of Wood and Wood Components
(1990)
R. A. Blanchette/A. R. Biggs (Eds.)
Defense Mechanisms of Woody Plants Against Fungi (1992)
S. Y. Lin/e. W. Dence (Eds.)
Methods in Lignin Chemistry (1992)
G. Torgovnikov
Dielectric Properties of Wood and Wood-Based Materials (1993)
F. H. Schweingruber
Trees and Wood in Dendrochronology (1993)
P. R. Larson
The Vascular Cambium: Development and Structure (1994)
M.-S. Ilvessalo-Pfaffli
Fiber Atlas: Identification of Papermaking Fibers (1994)
Philip R. Larson
The Vascular Cambium
Development and Structure
With 340 Figures
Springer-Verlag
Berlin Heidelberg New York London Paris
Tokyo Hong Kong Barcelona Budapest
Dr. PHILIP R. LARSON
Chief Plant Physiologist (Retired)
North Central Forest Experiment Station
USDA Forest Service
Rhinelander, WI 54501, USA
Series Editor:
T. E. TIMELL
State University of New York
College of Environmental
Science and Forestry
Syracuse, NY 13210, USA
Cover: Transverse section of Pinus lamberriana wood. Courtesy of Dr. Carl de Zeeuw,
SUNY College of Environmental Science and Forestry, Syracuse, New York
ISBN-13:978-3-642-78468-2 e-ISBN-13:978-3-642-78466-8
DOl: 10.1007/978-3-642-78466-8
Library of Congress Cataloging-in-Publication Data. Larson, Philip R. The vascular cambium: devel
opment and structure I Philip R. Larson. p. cm. - (Springer series in wood science) Includes
bibliographical references (p.) and index. ISBN-13:978-3-642-78468-2
I. Cambium. I. Title. II. Series. QK725.L29 1994 581.4'I-dc20 94-7963 CIP
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To my wife
Yvonne
Preface and Acknowledgements
A treatise on the vascular cambium must of necessity represent a compromise.
This necessity for compromise becomes obvious when one recognizes the per
vasiveness of the vascular cambium and the multiplicity of its derivative elements
as found in a large tree. No single volume can begin to adequately cover what is
known about this fascinating subject. The present volume, even though confined
to cambial development and structure, also represents a compromise. The decision
as to precisely what topics to include and how to discuss them was mine to make,
and I am solely responsible for the contents.
My interest in the cambium began many years ago as a graduate student at Yale
University. Beginning with early investigations of wood formation in trees, my re
search interests progressed through various stages of experimentation, finally
culminating in the structure/function relations of the cambium and its products.
These investigations, involving both anatomical and physiological studies, made
me even more acutely aware of the pervasiveness of the vascular cambium and the
difficulty of presenting it in a single volume. Not only is the vascular cambium the
progenitor of all xylem and phloem, but subsumed within these two broad cate
gories are numerous derivative anatomical elements of widely varying nature.
Moreover, both the cambium and its derivative elements are subject to structural
and functional modifications as the plant ages and its environment evolves.
Further contributing to the pervasiveness of the vascular cambium is its origin
from the procambium as I have stressed in the Introduction. When Professor
Timell first approached me to undertake the task of writing a book on the cam
bium, it was my intent to include the procambium. Interestingly enough, a com
prehensive coverage of the procambium-cambium relationship which places the
two parts of this continuum in proper perspective has never been attempted. None
theless, despite my strong interest in the subject, it soon became obvious that the
procambium and cambium comprised too large a topic for a single volume.
I have introduced my treatise with an historical review of the cambium quite
unlike those of other reviewers. Whereas others have highlighted specific events, I
have attempted to trace in considerable detail the tortuous pathways by which the
cambium eventually became recognized as a cellular, dividing tissue. The work of
Mirbel, who not only devoted his professional career to study of the cambium but
also contributed significantly to its elucidation, has been accorded special atten
tion. Here, and elsewhere in the text, the apology offered by De Bary (1877)
vm
might well apply: "From the older literature, I have perhaps cited too much for
many, and for others too little."
This historical approach has been adhered to in introducing other chapters and
sections in the text. It has been my experience over the years, both in speaking
with colleagues and in reading the literature, that most biologists profess know
ledge of the present status of a certain biological event or process but very few are
aware of how this knowledge was discovered. This oversight often leads to later
dubious claims of priority. As Esau (1969) has so aptly stated, "The pace of mo
dern research leaves little time for contemplative studies of historical documents."
Consequently, I have provided the historical background in an attempt to clarify
the priority for certain observations and discoveries.
In many cases, the original terminology has been retained when citing early pa
pers, not only to preserve the author's intended meaning but also to avoid injecting
unintended interpretations. For similar reasons, the reader might encounter appar
ent contradictions when comparing different sections of the text. This results in
part because of uncertainties and contradictions in the original papers and in part
because it was not possible to describe the subtle caveats, details, etc. brought
forth by each author.
Although I am the author of this book and bear sole responsibility for all errors
of omission and interpretation, lowe its existence to many others. I am indebted
to Professor Tore Timell, who not only suggested that I write the book, but also
offered valuable advice and encouragement during the years of preparation. Two
institutions provided invaluable services without which this work could never
have been completed. Personnel of the North Central Forest Experiment Station,
USDA Forest Service, my former employer, have graciously granted me office
space and full use of facilities and services since my retirement. I am particularly
indebted to Drs. R. D. Lindmark, Director of the North Central Forest Experiment
Station, St. Paul, Minnesota and J. G. Isebrands, Director's Representative and
Project Leader, Rhinelander, Wisconsin.
Among the invaluable services provided by the Forest Service was a coopera
tive arrangement between the Experiment Station and the University of Minnesota
Agricultural Library. Every reference cited in the text has been read either in the
original volume or a photoprint therefrom. The vast majority of these papers has
been provided by the dedicated librarians of the Experiment Station and the Uni
versity of Minnesota's Interlibrary Loan Department. Many additional papers have
been made available by the US Department of Agriculture Library in Beltsville,
Maryland.
My special thanks to those who offered invaluable comments and suggestions.
The manuscript was read in its entirety by Tore Timell, and selected chapters by J.
R. Barnett, G. P. Berlyn, B. G. Cumbie, R. A. Gregory, L. Kucera, C. M. Larson,
N. R. Lersten, C. H. A. Little, R. T. Riding, W. C. ShortIe, F. W. Taylor, F. W.
Telewski, E. A. Wheeler and B. F. Wilson.
Contents
1 Introduction
2 Fossil Evidence 7
3 Historical 9
3.1 Derivation of the Word Cambium 9
3.2 The Concept of Cambium as a Tissue 10
3.2.1 The Role of Cell Division 24
3.2.2 The Role of Cell Protoplasm 30
4 Defining the Cambium 33
4.1 Cambial Cell Theories 33
4.1.1 "Siamese Twin" Theory of Hartig 34
4.1.2 Uniseriate Initial Theory of Sanio 34
4.1.3 Multiseriate Theory of Raatz 42
4.1.4 Uniseriate vs. Multiseriate Theories 49
4.2 Special Criteria 58
4.2.1 Sanio's Rules 58
4.2.2 Additional Rules 59
4.2.3 Role of Mother Cells 60
4.2.4 Tangential Cell Alignment 61
4.3 Reconciliation 63
4.4 Terminology 66
4.5 Special Features 68
4.5.1 Sanio's Four 68
4.5.1.1 Xylem 68
4.5.1.2 Phloem 80
4.5.2 Bars of Sanio -Trabeculae 84
4.5.3 Rims of Sanio -Crassulae 92
4.5.4 Bars vs. Rims of Sanio 95
4.5.5 Monogene vs. Polygene Cambia 95
4.5.6 Cambial vs. Apical Initials 96
X
5 Cambial Cell Characteristics 99
5.1 Cell Shape 99
5.1.1 Effect of Cell Shape on Division 103
5.2 Fusiform Initial Size 105
5.2.1 Fusiform Initial Volume 108
5.3 Ultrastructure 108
5.3.1 Nucleus 110
5.3.1.1 Fusiform Initials 110
5.3.1.2 Ray Cell Initials 116
5.3.1.3 Nuclear Migration 119
5.3.1.4 DNA Content 119
5.3.1.5 Nucleoli 122
5.3.2 The Vacuole 124
5.3.2.1 Protoplasmic Streaming 126
5.3.3 Organelles 128
5.3.3.1 Gymnosperms 129
5.3.3.2 Dicotyledons 130
5.4 Cytochemistry 133
5.4.1 Starch 133
5.4.1.1 Gymnosperms 134
5.4.1.2 Dicotyledons 134
5.4.2 Lipids 135
5.4.3 Proteins 136
5.4.4 Phytoferritin 137
5.4.5 Tannins 139
5.4.6 Chlorophyll 139
5.4.7 Miscellaneous Constituents 140
5.5 Cambial Cell Division 140
5.5.1 Mitotic Index 146
5.5.2 Cambial Cell Cycle 148
5.6 Cambial Aging 152
6 Anticlinal Cambial Divisions 155
6.1 Pseudotransverse and Transverse Divisions 155
6.1.1 Occurrence 155
6.1.1.1 Analysis 160
6.1.2 Cell Length at Division 162
6.1.3 Cell Wall Orientation 167
6.1.4 Length of Cell Plate 174
6.1.5 Cell Elongation 178
6.1.5.1 Proximal End Growth 178
6.1.5.2 Distal End Growth 180
6.1.5.3 Sister Cell Polarity 183
6.2 Radial Anticlinal Divisions 186
XI
6.2.1 Storied Cambia 186
6.2.1.1 Storied Rays 196
6.3 Fates of Initial Cells 199
6.3.1 Survival of Initial Cells 199
6.3.1.1 Fusiform Cell Length 199
6.3.1.2 Ray Contacts 201
6.3.2 Failure of Initial Cells 203
6.3.2.1 Cell Elimination 204
6.3.2.2 Segmentation to Parenchyma 209
6.3.2.3 Segmentation to Rays 214
6.3.2.4 Complete Segmentation 216
6.3.2.5 Partial Segmentation 221
6.4 Tangential Increase of Cambium 227
6.4.1 Nonstoried Cambia 228
6.4.1.1 Fusiform Initials 228
6.4.1.2 Ray Cell Initials 235
6.4.2 Storied Cambia 236
6.5 Frequency of Anticlinal Divisions 237
6.5.1 Age of Tree 238
6.5.2 Rate of Growth 240
6.5.2.1 Eccentric Stems and Branches 245
6.5.3 Location Within the Growth Ring 247
6.5.4 Position in the Tree 250
6.5.4.1 Height in the Stem 250
6.5.4.2 Branches 251
6.5.4.3 Branch Crotches 252
6.5.4.4 Roots 256
6.5.4.5 Living Stumps 258
6.5.5 Variations Among Species 260
6.5.6 Biennial Frequency 261
6.6 Aberrant Anticlinal Divisions 261
6.6.1 Localized Domains 261
6.6.2 Migrating Domains 265
6.6.2.1 Wavy Grain 269
6.6.2.2 Interlocked Grain 272
6.6.2.3 Spiral Grain 279
6.6.2.4 Whirled Grain 283
6.6.2.5 Circular Vessels and Tracheids 285
6.6.3 Wave Pattern of Domains 288
6.6.4 Theoretical Interpretations 290
6.7 Mechanism of Cell Elongation 293
6.7.1 Sliding Growth 293
6.7.2 Symplastic Growth 297
6.7.3 Intrusive and Interpositional Growth 299