Comparative Wood Anatomy: Systematic, Ecological, and Evolutionary Aspects of Dicotyledon Wood

447 Pages·1988·19.27 MB·English

by  Dr. Sherwin Carlquist (auth.)

Springer Series in W ood 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. Areher
Growth StressesandStrains in Trees (1986)
W. E. Hillis
Heartwood and Tree Exudates (1987)
S. Carlquist
Comparative Wood Anatomy (1988)
L. W. Roberts/P. B. GahanfR. Aloni
Vascular Differentiation· and Plant Growth Regulators (1 988)
Sherwin Carlquist
Comparative
Wood Anatomy
Systematic, Ecological, and
Evolutionary Aspects of Dicotyledon Wood
With 101 Figures
Springer-Verlag
Berlin Heidelberg GmbH
Dr. SHERWIN CARLQUIST
Rancho Santa Ana Botanic Garden
and Pomona College
Claremont, CA 91711-3101, USA
Series Editor:
T. E. TIMELL
State U niversity of N ew Y ork
College of Environmental
Science and Forestry
Syracuse, NY 13210, USA
Cover: Transverse section of Pinus lambertiana wood- Courtesy of Dr. Carl de Zeeuw,
SUNY College of Environmental Science and Forestry, Syracuse, New York
ISBN 978-3-662-21716-0 ISBN 978-3-662-21714-6 ( eBook)
DOI 10.1007/978-3-662-21714-6
Library of Congress Cataloging-in-Publication Data.
Carlquist, Sherwin John, 1930-Comparative wood anatomy. (Springer series in wood science)
Bibliography: p. 358 Includes index. 1. Wood-Anatomy. 2. Dicotyledons. I. Title. II. Series.
QK647.C37 1988 583'.041 88-6465
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© Springer-Verlag Berlin Beideiberg 1988
Originally published by Springer-Verlag Berlin Beideiberg New York in 1988
Softcover reprint of the hardcover 1st edition 1988
The use of registered names, trademarks, etc. in this publication does not imply, even in the
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2131/3130-543210
Preface and Acknowledgments
One reason for presentation of a book on comparative wood anatomy
is that no recent book solely on this topic has been written. Those
familiar with wood anatomy are certainly familiar with other books
dealing with various ofits aspects, such as identification, dendrology,
dendrochronoloGy, and other aspects closely related to comparative
wood anatomy. These books are certainly part of the field of
comparative wood anatomy and are cited in Chapter 1 and elsewhere
in this book. The purpose oftbis book is not to duplicate those efforts,
but to offer a volume that has a different focus. This book assumes
that the readerwill have an elementary knowledge ofthe subject, such
as might be obtained in a course in plant anatomy.
Because comparative wood anatomy deals with descriptions of
wood features, an attempt has been made to cover as many wood
features as possible that vary with relation to the taxonomic system.
Each worker may have a different way of producing a description,
but with a reason. The present compilation attempts to see the value in
all of these variants and take into account the collective wisdom
in wood descriptions. The nature of that wisdom is continually in
the process of change, and this book follows significant changes that
have occurred in wood anatomy during the past several decades.
The work of I. W. Bailey and his students gave an evolutionary
dimension to comparative wood anatomy. Bailey realized that
adaptation to ecology is a central theme in wood evolution, and so
the field of ecological wood anatomy (which may be considered
functional wood anatomy on a comparative basis) becomes an
integral part of comparative studies. In describing differences among
species or genera, we are usually describing differences in adaptation
to ecological features.
The synthesis between descriptive, functional, ecological, and
physiological aspects of wood anatomy has been slow and difficult.
This synthesis is too broad for any one person to effect, so various
workers must contribute to it. The data from comparative wood
anatomy appeal to me as vital sources of hypotheses - and of
materials for testing ofhypotheses. Comparative workers do not need
to feel inferior to those who do experimental work, because natural
experiments in ecological wood anatomy have produced compelling
patterns, and those interested in wood physiology can ignore those
patterns only at their peril. Large genera, distributed into different
habitats, ·a re like replicates of an experimental material, with the
additional advantage of time for selection of optimal wood plans for
VI Preface
each ecological habitat (we must note that wood is not the sole tool
whereby a plant deals with water economy).
However, comparative workers should be vitally interested in the
results of wood physiology, because this field can show with clarity
and precision exactly how wood adaptations work. Thus a bridge
between the two fields is in the process of construction.
The transitional nature ofthe field of comparative wood anatomy
and the enormity of Iiterature in this area make construction of a
book difficult. No two individuals would have selected material
similarly; my selection, although idiosyncratic, hopefully can serve a
range of interests. No book is a substitute for the primary Iiterature
of a field, especially those with extensive citation of papers.
I take pleasure in acknowledging the many who have aided my
studies in wood over the years. Selecting a few is difficult, for I have
leamed directly, and by letters, from many, including Peter Baas,
I. W. Bailey, C. R. Metcalfe, Regis Miller, Rudolf Schmid, William
L. Stern, Elizabeth Wheeler, Martin Zimmermann, and a large num
ber of knowledgeable persons who have reviewed manuscripts of my
papers. I am grateful to Dr. Thomas S. Elias for his practical role as
Director ofthe Rancho Santa Ana Botanic Garden, site ofmy work,
and to Dr. Robert F. Thome for his systematic ideas. Among those
who have fumished me with material, I must cite Dr. Regis B. Miller
and Dr. Peter H. Raven above all-but many others deserve mention
as given in my papers on various taxonomic groups and wood
topics. Students and former students who have provided valuable
assistance, useful interaction, and a source of inspiration include
Jonathan R. Davis, Vincent M. Eckhart, David A. Hoekman, Gary
M. W allace, David Wheat, and Scott Zona.
Claremont, USA s. CARLQUIST
Spring 1988
Contents
1 Methods for Comparative Wood Anatomy Studies . . . 1
1.1 Techniques for Comparative W ood Anatomy Studies 1
1.2 Sources of Wood Sampiesand Wood SampiePreservation 3
1.3 Literature on Comparative Anatomy . . . . 5
1.4 Terminology . . . . . . . . . . . . . . 6
1.5 Sequence of Features in Wood Descriptions. 7
1.6 Magnifications 11
2 Growth Rings . 12
2.1 Introduction . 12
2.2 Factors Affecting Growth-Ring Formation 12
2.3 Factors in Classification of Growth Rings 13
2.4 Description of Growth Ring Types 17
2.5 Interxylary Cork . . . . . . 36
2.6 Modifiability of Growth Rings . . 37
2. 7 Experimental Studies . . . . . . 37
2.8 Cell Length with Respect to Growth Rings 38
2.9 Systematic Distribution of Growth Rings . 38
3 Vessel Elements . . . . . 40
3.1 Definition . . . . . . . 40
3.2 Types of Vessel Elements. 41
3.3 Vessel Dimensions. 43
3.4 Vessel Grouping 46
3.5 Vessel Density . . 54
3.6 Origin of Vessels . 54
3.7 The Perforation Plate 61
3.8 Lateral Wall Pitting of Vessels 73
3.9 Crateriform Pits . . . . . 83
3.10 Vesturing . . . . . . . . . 84
3.11 Verrucae on Vessel Walls. . . 89
3.12 Helical Sculpture on Vessel Walls 89
3.13 Vessel Wall in Transection 99
3.14 Tyloses . . 102
3.15 Trabeculae . . . . . . . 103
4 Imperforate Tracheary Elements 104
4.1 Definitions and Terminology . 104
VIII Contents
4.2 Nature of the Bordered Pit . . . . . . . . . 107
4.3 Evolution from Tracheids to Libriform Fibers. 108
4.4 Tracheids of Vessel-less Dicotyledons . . . . 111
4.5 True Tracheids in Vessel-Hearing Dicotyledons 116
4.6 Fiber-Tracheids . . . . . . . . 120
4.7 Libriform Fibers . . . . . . . 124
4.8 Septate Fibers and Living Fibers 129
4.9 Fiber Dimorphism. . . . . . 132
4.10 Fiber Distribution Anomalies . 133
4.11 Vascular Tracheids . 133
4.12 Vasicentric Tracheids 135
4.13 Reaction Wood . . . 144
4.14 Trabeculae . . . . . 147
4.15 Contents oflmperforate Tracheary Elements 148
4.16 Excluded Types oflmperforate Tracheary Elements 148
5 Axial Parenchyma . . . . . . . . . . 150
5.1 Definition, Means of Observation . . . 150
5.2 Types of Axial Parenchyma Distribution 151
5.3 Parenchymatization . . . . . . . . . 165
5.4 Parenchyma Proliferation. . . . . . . 168
5.5 Histological Details of Axial Parenchyma. 168
5.6 Pith Flecks . 172
6 Rays 174
6.1 Terminology, Definitions . 174
6.2 Ray Dimensions . . . . 175
6.3 Ray Types: Problems in Definition 178
6.4 The Kribs Ray Types . . 181
6.5 Paedomorphic Ray Types 190
6.6 Raylessness. . . 194
6. 7 Aggregate Rays . . . 199
6.8 Ray Dimorphism . . 201
6.9 Interconnected Rays . 202
6.10 Intermittent Rays 202
6.11 Other Ray Types . . 202
6.12 Ray Histology . . . 202
6.13 Systematic Distribution of Ray Types 213
7 Cell Contents, Secretory Structures . 216
7.1 Crystals . 216
7.2 Cystoliths . . . . . . . . . . . 238
7.3 Starch ............ . 240
7.4 Silica Bodies and Other Silica Deposits . 241
7.5 Laticifers and Latex Deposits . 245
7.6 Tanniniferous Tubules . . . . . . . . 247
Contents IX
7. 7 Oil and M ucilage Cells . . . . . . . 24 7
7.8 Cells with Amorphous Deposits . . . 251
7.9 Intercellular Canals (Secretory Canals) 251
8 Cambial V ariants (Anomalous Secondary Growth) 256
8.1 Classification and Nature of Types . . . . . . 256
8.2 Successive Cambia; the Primary Thickening Meristem
and its Products . . . . . . . . . . . . . . . . 257
8.3 Interxylary Phloem Produced by a Single Cambium 262
8.4 Cambia Normal in Products, but Abnormal in
Conformation or Dispersion . . . . . . . . . . 266
8.5 Combinations of Cambial Variants . . . . . . . 273
8.6 Phylogeny, Habit, and Ecology of Cambial Variants 274
9 Cambial Ontogeny (Storying, Paedomorphosis Other
Changes) . . . . . . . . . . . . . . . . . . . . . 278
9.1 Introduction . . . . . . . . . . . . . . . . . . . 278
9.2 Increase in Circumference of the Cambium and Storying 278
9.3 Ray Ontogeny . . . . . . . . . . . . . . .... 288
9.4 Paedomorphosis . . . . . . . . . . . . . . . . . 288
9.5 Other Wood Phenomena Related to Cambial Ontogeny 295
10 Systematic Application of Wood Data . 297
10.1 Introduction . . . . . . . . . . . 297
10.2 Types of Systematic Applications . . 298
10.3 Systematic Value ofWood Features . 305
10.4 Formats for Reporting Taxonomie Features of Woods 312
11 Evolution in Wood: An Ecological/Functional Synthesis 315
11.1 Introduction . . . . . . . . . . . . . . . . . . 315
11.2 I. W. Bailey and the "Major Trends ofXylem Evolution" 315
11.3 Irreversibility . . . . . . . . . . . . . . . . 329
11.4 Features Related to the Major Trends of Xylem
Evolutions, but with Modifications . . . . . . 332
11.5 Features Not Related to the Major Trends of Xylem
Evolution: Conductive Efficiency and Safety . . . . 335
11.6 Features Not Related to the Major Trends of Xylem
Evolution: Features Related to Photosynthates . . . 340
11.7 Features Not Related to the Major Trends of Xylem
Evolution: Mechanical Strength . . . . . . . . . . 341
11.8 Features Not Related to the Major Trends of Xylem
Evolution: Water Storage. . . . . . . . . . . . . 343
11.9 Features Not Related to the Major Trends of Xylem
Evolution: Predation Deterrence . . . . . . . . . . 344