Plant Structure: Function and Development: A Treatise on Anatomy and Vegetative Development, with Special Reference to Woody Plants

531 Pages·1993·29.36 MB·English

by  Dr. John A. Romberger

John A. Romberger · Jane F. Hill
Zygmunt Hejnowicz
Plant Structure:
Function and
Development
A Treatise on Anatomy and
Vegetative Development, with
Special Reference to Woody Plants

J. A. Romberger· Z. Hejnowicz . 1. F. Hill
Plant Structure:
Function and Development
A Treatise on Anatomy and
Vegetative Development,
with Special Reference to Woody Plants
With 57 Figures
Springer-Verlag
Berlin Heidelberg GmbH
DR. JOHN A. ROMBERGER (Emeritus Collaborator)
US Department of Agriculture
Beltsville Agricultural Research Center - West
Climate Stress Laboratory, Bldg. 046A
10300 Baltimore Avenue, Beltsville, Maryland 20705, USA
and: 2005 Forest Hill Drive
Silver Spring, Maryland 20903, USA
PROF. ZYGMUNT HEJNOWICZ
Department of Biophysics and Cell Biology
Silesian University
ul. Jagiellonska 28
40-032 Katowice, Poland
and: Botanical Institute
University of Bonn
Venusbergweg 22
D-5300 Bonn 1, FRO
DR. JANE F. HILL (Volunteer Collaborator)
US Department of Agriculture
Beltsville Agricultural Research Center - West
Climate Stress Laboratory, Bldg. 046A
10300 Baltimore Avenue, Beltsville, Maryland 20705, USA
and: 8211 Hawthorne Road
Bethesda, Maryland 20817, USA
ISBN 978-3-662-01664-0 ISBN 978-3-662-01662-6 (eBook)
DOI 10.1007/978-3-662-01662-6
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© Springer-Verlag Berlin Heidelberg 1993
Originally published by Springer-Verlag Berlin Heidelberg New York in 1993
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1YPesetting: Camera ready by senior author
Production Editor: Renate Miinzenmayer
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Preface
This book is about the developmental anatomy of large, complex plants, particularly
of the woody plants that grow and survive for decades or centuries. It is focused on
the meaning of that anatomy, that integrated structure, as a determinant of effective
function. A pervading theme is that the plant structures that have survived "selection"
processes during the eons of organismal evolution, within the larger context of geo
logic and climatic evolution, are well attuned to biochemical and biophysical principles
that determine and define efficient function.
The sets of structure-and-function couples existing in the various plant taxa differ
so widely that generalities are often difficult to discern. This diversity is due partly to
the broad range of ecological conditions to which higher plant organisms have become
adapted under stresses imposed by competition and continual climatic change. It is also
due to the tendency of different taxa, with their different complements of inherited
information, to respond to similar situations in different ways. Cognizant of this
reality, we have tried throughout the book to avoid generalizing too broadly on the
basis of data from the relatively small fraction of plant species that have as yet been
studied.
This book is intended for those who have already studied the anatomy and develop
ment of plants. It is addressed to advanced students, teachers, and researchers in the
interrelated fields of botany, forestry, horticulture, and agronomy, and to others having
professional interests in the culture of woody plants and the stewardship of ecosystems.
The book is also addressed to those, of whatever broader discipline, who invoke
theoretical biology in efforts to understand how the higher plant, as a complex organ
ism, copes with its biotic and abiotic environment, grows, develops, and reproduces
its kind - while also supporting the animal world (including us). Especially it is
addressed to those who, by study and research, seek to narrow the wide gap between
the cellular and molecular biology approaches to understanding the format and content
of inherited information, and the actual morphogenesis and integrated functioning of
higher plant organisms. We feel that presuming to study or practice the various
botanical subdisciplines without first gaining a full appreciation of the meanings
inherent in developmental plant anatomy is somewhat like attempting to practice
medicine without first having acquired an understanding of the anatomy, development,
and functioning of the normal human organism.
vi Preface
By citation of a range of significant references, including some less recent ones, we
have sought to point out access routes to the origins of ideas, to the rich research
traditions, and the stores of basic information in the world literature -lest we become
too strongly influenced by recent thinking and work still to be thoroughly evaluated by
broad scientific opinion as it evolves over time.
We hope that the ideas and interpretations embodied in this volume will serve as
stimulants to those planning and conducting research on plant growth and development,
and also that the book will be useful as a guide to the thinking and literature pertinent
to advanced courses on plant structure and development; for some it may, perhaps,
serve as a textbook.
Though we initially had broader aspirations, this book is focused on vegetative
growth and development. Limitations of space precluded a treatment of reproductive
development, and of morphogenesis in fruits and seeds. We have, however, included
a chapter on embryogeny as the beginning of development of the individual higher
plant organism.
October, 1992 The Authors
Acknowledgements
The authors express their appreciation of the encouragement and moral support given
by their various present and former colleagues in Poland, Germany, and the United
States during preparation of this book. The senior author (JAR) gratefully acknow
ledges support from the Interacademy Exchange Program between the counterpart
organizations in the USA and Poland, which enabled him to work at Silesian Univers
ity in Katowice with the second author (ZH) in 1981 and 1983. He also acknowledges
the gracious hospitality extended to him by that institution. This book was planned and
partially drafted during those visits. The senior author further gratefully acknowledges
the support and hospitality received from The Institute of Agricultural Sciences,
College of Agriculture (A. R. Lublin) in Zamosi, Poland, during a visit in 1985.
Work on this book was continued both jointly (by ZH and JAR) during such visits, and
separately as conditions allowed.
Two of us (JAR and JFH), both having been affiliated with the former Forest
Physiology Laboratory of the US Department of Agriculture, Forest Service, in
Beltsville, Maryland, are pleased to acknowledge the hospitality and professional
recognition afforded by appointments as Volunteer Collaborators at the Climate Stress
Laboratory of the US Department of Agriculture, Agricultural Research Service, in
Beltsville, Maryland. We also acknowledge, with sincere appreciation, the splendid
cooperation and assistance provided to us over the years by the staff members of The
National Agricultural Library in Beltsville. Their help was truly indispensable.
The second author (ZH) expresses his appreciation to the administration of
Silesian University for enabling him to accept a Visiting Professorship at the University
of Bonn, and to the latter institution for allowing him to continue working part-time
on this book while in residence there. The senior author (JAR) also expresses his
appreciation of hospitality and working space provided to him by the University of
Bonn during visits in 1983 and 1988.
The authors collectively express their indebtedness to Dorota Hejnowicz and
to Karen L. Parker for their dedicated work in skillfully drawing, or redrawing, the
technically accurate illustrations that the book required. We also acknowledge the
invaluable help of many of our associates who consented to read critically, review, or
proofread the entire manuscript or parts of it. We want especially to mention the
contributions of Robert D. Warmbrodt, William Van Der Woode, Claude L. Brown,
Robert McGaughy, George C. Martin, and Kitren G. Weis. The senior author also
viii Acknowledgments
acknowledges the genuinely valuable proofreading done by his wife, Margery D.
Romberger.
The authors gratefully acknowledge the pennissions to redraw or reproduce illustra
tions, from copyrighted publications, granted by the holders of those copyrights. We
are pleased specifically to mention: Panstwowe Wydawnictwo Naukowe, Warsaw
(publisher of Hejnowicz 1973,1980); Wiley and Sons, New York (publisher of Esau
1965a); Wissenschaftliche Veriagsgesellschaft mBH, Stuttgart (publisher of Braun
1963); SIR Publishing, Wellington (publisher of New Zealand Journal of Botany);
Blackwell Scientific Publications, Oxford (publisher of Clowes 1961); McGraw-Hill
Publishing Co., New York (publisher of Eames and MacDaniels 1947); Springer
Verlag (Wien), Vienna (publisher of Protoplasma); Botanical Society of America,
Ames (publisher of American Journal of Botany); Cambridge University Press, New
York (publisher ofWilIiams 1975); Cambridge University Press, Sheffield (publisher
of New Phytologist); University of Chicago Press (publisher of Botanical Gazette);
Carl Winter Universitiitsverlag, GmbH, Heidelberg (publisher of Magdefrau 1951);
Forest Products Research Society, Madison (publisher of Wood Science); Society of
Wood Science and Technology, Madison (publisher of Wood and Fiber); and The
National Research Council of Canada, Ottawa (publisher of Canadian Journal of
Botany). Details pertaining to sources of specific illustrations are given in the figure
captions and in the References.
Contents
Part I. Functional Anatomy
1 Introduction
1.1 The Subject ........................................ 3
1.2 Hierarchy: A Concept of Ordered Relations. . . . . . . . . . . . . . . . . . .. 5
1.3 Geometry and Dimensions of Cells ......................... 6
1.4 Cell Walls ......................................... 7
1.4.1 General Aspects. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7
1.4.2 Pits ................................. . . . . .. 8
1.4.3 Cell-Wall Polymers. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10
1.4.4 Water in Cell Walls. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11
1.4.5 Genesis and Ultrastructure of Cell Walls ............... 12
1.4.6 Incrustation of Cell Walls . . . . . . . . . . . . . . . . . . . . . . . .. 16
1.4.7 Adcrustation of Cell Walls ........................ 16
1.5 The Duality of Spatial Systems Within Plants . . . . . . . . . . . . . . . . . .. 17
1.5.1 The Symplasm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18
1.5.2 The Apoplasm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19
1.5.3 Transfer Between Apoplasm and Symplasm . . . . . . . . . . . .. 20
1.6 The Morphological Classification of Tissues ................... 20
2 Protective Systems - Boundaries
2.1 Introduction ........................................ 23
2.2 Dermal Tissues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 23
2.2.1 Shoot Epidermis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24
2.2.1.1 Cuticle and Epicuticular Waxes. . . . . . . . . . . . . .. 25
2.2.1.2 Cuticle and Water Relations . . . . . . . . . . . . . . . .. 27
2.2.2 Rhizodermis ............................... . 28
2.3 Exodermis, Endodermis, and Metacutis ..................... . 29
2.4 Periderm: Phellogen, Phellem, Phelloderm ................... . 30
2.5 Rhytidome, or Outer Bark .............................. . 31
3 Absorbing Systems
3.1 General Perspectives .................................. 33
X Contents
3.2 Structural Aspects of Absorption from Outside the Plant. . . . . . . . . . .. 34
3.2.1 Embryonal and Cotyledonary Adaptations .............. 34
3.2.2 Shoot Tissues and Adaptations: Transcuticular Absorption. . .. 35
3.2.3 Root Tissues and Adaptations ...................... 36
3.2.3.1 Rhizodermis and Root Hairs. . . . . . . . . . . . . . . .. 36
3.2.3.2 Exodermis and Cortex .................... 38
3.2.3.3 Mycorrhizal and Cluster (Proteoid) Roots ........ 38
3.2.3.4 Other Adaptations ....................... 41
3.3 Structural Aspects of Internal Absorption and Transfer. . . . . . . . . . . .. 42
3.3.1 Wall Ingrowths and Transfer Cells . . . . . . . . . . . . . . . . . .. 42
3.3.2 Plasmatubules ................................ 44
4 Supportive Systems
4.1 Introduction ........ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45
4.2 Organs, Tissues, and Cells as Mechanical Components ............ 45
4.3 Mechanics of Complex Beams: General Consideration ........... " 47
4.3.1 Normal Stresses, Normal Strains, and the Modulus of Elas-
ticity ..................................... 47
4.3.2 Bending and Stiffness ........................... 48
4.3.3 Shear Stresses and Strains ....................... " 50
4.3.4 Buckling and the Slenderness Ratio. . . . . . . . . . . . . . . . . .. 51
4.4 Applications of Mechanics to Parenchyma Cells and Tissues ......... 53
4.4.1 Cell-Wall Tensions and Parenchyma-Tissue Strength ....... 53
4.4.2 Arrangement of Reinforcing in Cell Walls .............. 55
4.5 Dynamic Loading and Organ Resilience .... . . . . . . . . . . . . . . . . .. 57
4.5.1 Strain Energy ............................... " 57
4.5.2 Torsions ................................... 59
4.6 Growth-Induced Tissue Stresses ........................... 60
4.7 Design and Construction Strategies ......................... 62
4.7.1 Cantilever Beams and Stem Taper ................... 62
4.7.2 Design for Resistance to Buckling ................... 63
4.7.3 Design for Simultaneous Resistance to Static and Dynamic
Loads .................................. " 64
5 Photosynthetic System
5.1 Structural Aspects of Photosynthetic Function .................. 67
5.2 Alternate Pathways of Carbon Fixation . . . . . . . . . . . . . . . ...... " 68
5.3 Structural Implications of Photosynthetic Pathways ............... 71
5.3.1 Leaf Structure in C3 Plants. . . . . . . . . . . . . . . . . . . . . . .. 72
5.3.2 Leaf Structure in C4 Plants. . . . . . . . . . . . . . . . . . . . . . .. 74
5.4 CAM and Other Variations .............................. 76
5.5 Leaf Structure in Relation to Optical Properties ................. 77
6 Storage Systems
6.1 Storage and Survival. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79
6.2 General Characteristics of Storage Organs and Tissues .............. 80