Table Of ContentCRC
Handbook
of
Lichenology
Volum e III
Editor
Margalith Galun, Ph.D.
Professor
Department of Botany
The George S. Wise Faculty of Life Sciences
Tel Aviv University
Tel Aviv, Israel
Boca Raton London New York
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PREFACE
Lichens are among the most widely distributed eukaryotic organisms in the world. There
are, to date, about 13,500 lichen species known, which accounts for approximately 20% of
all the fungi described. All are the result of a symbiotic association between two unrelated
organisms — a fungus and an alga (or cyanobacterium) — which, when fully integrated,
form a new biological entity with very little resemblance to either one of its components.
Despite their wide ecological amplitude and abundance, also in extreme environments
where often other plants cannot exist, lichens have received relatively little attention by plant
physiologists. It was not until 1984 that the first international conference on lichen physiology
and cell biology was held (organized by D. H. Brown, in Bristol). The second conference,
devoted in part to lichen physiology, took place in 1986 (organized by E. Peveling, in
Munster).
I have attempted to invite as many as possible lichenologists who have made significant
contributions to lichen research, to contribute chapters to this book. Each author has been
encouraged to approach his subject in his own way, in order to communicate the author’s
enthusiam to the reader. Naturally, in a book where each chapter deals with its own topic,
there is bound to be some overlap of subject matter. Thus, people who are interested in a
specific subject and read only one of the chapters, need to have some background pertinent
to that subject. The reader will also find some contradictory statements in the various chapters,
mainly on assumptive aspects, which I did not find necessary to unify, in order to stimulate
further investigations.
Topics which have not been comprehensively surveyed in other treatises or are scattered
in different journals, such as Nitrogen Metabolism (Chapter VI.B), Enzymology (Chapter
VI.C), and Ecophysiological Relationships in Different Climatic Regions (Chapter VII.B.2),
are here given more substantial scope. The Algal Partner (Chapter II.B) is much more
extended that The Fungal Partner (Chapter II. A), because the systematic status of the pho-
tobionts has not been treated before in an integrated manner.
The terms “phycobiont” for the algal symbiont and “mycobiont” for the fungal symbiont
of lichens have been introduced by G. S. Scott in 1957 [Lichen Terminology, Nature
(London), 179,486, 1957]. After the “blue-green algae’’ were classified as “cyanobacteria’’
it was proposed (Ahmadjian, V., Holobionts Have More Parts, International Lichenological
Newsletter, 15, 19, 1982) that “photobiont’’ should be used for the photosynthetic partner(s)
of the lichens and more specifically “cyanobiont’’ for the cyanobacterial symbionts and
“phycobiont’’ for the eukaryotic algal symbionts. This terminology has been adopted here,
except for Chapter II.B (Tschermak-Woess) where, upon the author’s request, the previous
terms “phycobiont’’ and blue-green algae (Cyanophyceae) appear.
The names of the lichens cited in the various chapters are always those of the original
articles. This may, in many cases, not coincide with the modernized system by Haffelner
(Chapter X).
It is my hope that this book will bring lichenology to the notice of biology researchers ,
teachers, and students, as an important phenomenon in the mainstream of biology.
I greatly appreciate the efforts of the contributors and am especially indebted to Dr. Paul
Bubrick, who encouraged me to launch this endeavor and who was very helpful in the
editing of several chapters. I also wish to thank Dr. Leslie Jacobson, who read and commented
on the chapters I contributed to this book, and Ms. Ruth Direktor and Ms. Henriette van
Praag for their skillful secretarial assistance.
Finally, I am deeply grateful for the help, forbearance, and encouragement of my family.
Margalith Galun
THE EDITOR
Margalith Galun, Ph.D. is Professor of Botany at The George S. Wise Faculty of Life
Sciences of Tel Aviv University. She received her M.Sc. and Ph.D. degrees from The
Hebrew University of Jerusalem. Dr. Galun is a member of several professional and scientific
organizations and in some she holds executive positions, such as Vice-President of the
International Association for Lichenology, while in others she serves on the Executive
Committee, such as that of the International Mycological Association and of the Israel
National Collections of Natural History. She is the editor-in-chief of the journal Symbiosis
and is a member of the editorial board of the journals Israel Journal of Botany, Endocytobiosis
and Cell Research, and Lichen Physiology and Biochemistry. Dr. Galun has been the recipient
of many research grants and is the author or co-author of two books and of about 90 research
articles, including reviews, symposia, and chapters in books. Her current major research
interests relate to the interaction between symbionts of plant symbiotic systems.
CONTRIBUTORS
Volume III
Madalena Baron, M.Sc. Marcello Iacomini, Ph.D.
Professor Professor
Departamento de Bioquimica Departamento de Bioquimica
Universidade Federal do Parana Universidade Federal do Parana
Curitiba, Parana, Brazil Curitiba, Parana, Brazil
Adiva Shomer-Ilan, Ph.D.
Paul Bubrick, Ph.D. Senior Lecturer
Research Associate Department of Botany
Department of Plant Pathology The George S. Wise Faculty of Life
University of California Sciences
Riverside, California Tel-Aviv University
Tel-Aviv, Israel
Bazyli Czeczuga, Sc.D.
John L. Innes, Ph.D.
Professor
Research Scientist
Department of General Biology
Forestry Commission
Medical Academy
Alice Holt Lodge
Bialystok, Poland
Wrecclesham
Famham, Surrey, England
Margalith Galun, Ph.D.
David Jones, Ph.D.
Professor
Principal Scientific Officer
Department of Botany
Department of Microbiology
The George S. Wise Faculty of Life
Macaulay Land Use Research Institute
Sciences
Craigebuckler, Aberdeen, Scotland
Tel-Aviv University
Tel-Aviv, Israel
David H. S. Richardson, D.Phil.
Professor
Philip A. J. Gorin, Ph.D. School of Botany
Professor University of Dublin
Departamento de Bioquimica Trinity College
Universidade Federal do Parana Dublin, Ireland
Curtibiba, Parana, Brazil
Reuven Ronen, Ph.D.
Research Associate
Josef Hafellner, Ph.D. Department of Botany
Universitats Dozent The George S. Wise Faculty of Life
Department of Botany Sciences
Karl Franzens University Tel-Aviv University
Graz, Austria Tel-Aviv, Israel
TABLE OF CONTENTS
Volume I
SECTION I: INTRODUCTION
I. The True Nature Lichens — A Historical Survey...............................................................3
Jacob Lorch
SECTION II: LICHEN COMPONENTS
II. A. The Fungal Partner...........................................................................................................35
David L. Hawksworth
II. B. The Algal Partner...........................................................................................................39
Elisabeth Tschermak-Woess
SECTION III: THE LICHEN THALLUS
III. The Lichen Thallus................................................................................................................95
Hans Martin Jahns
SECTION IV: THE FUNGUS-ALGA RELATION
IV. The Fungus-Alga Relation.................................................................................................147
Margalith Galun
SECTION V: REPRODUCTION
V. A. Asci, Ascospores, and Ascomata..............................................................................161
Andre Bellemere and Marie Agnes Letrouit-Galinou
V. B. Conidiomata, Conidiogenesis, and Conidia..............................................................181
David L. Hawksworth
SECTION VI: LICHEN PHYSIOLOGY
VI. A. Carbon Metabolism.....................................................................................................197
Margalith Galun
VI.B. Nitrogen Metabolism.....................................................................................................201
Amar Nath Rai
VI.C. Lichen Enzymology.......................................................................................................239
Carlos Vicente and Marfa Estrella Legaz
INDEX 287
TABLE OF CONTENTS
Volume II
SECTION VII: LICHEN PHYSIOLOGY AND ECOPHYSIOLOGY
VILA. Ecology
VII.A.l. Substrate Colonization, Growth, and Competition.................................................3
Richard A. Armstrong
VII.B. Ecophysiology
VII.B.l. Water Relations.........................................................................................................17
Philip W. Rundel
VII.B.2. Ecophysiological Relationships in Different Climatic Regions...........................37
Ludger Kappen
VII.B.3. Physiological Buffering...........................................................................................101
John F. Farrar
VII.C. Contribution of Lichens to Ecosystems....................................................................107
Mark R. D. Seaward
SECTION VIII: CELLULAR INTERACTIONS
VIII.A. Effects of Symbiosis on the Photobiont..................................................................133
Paul Bubrick
VIII.B. Effects of Symbiosis on the Mycobiont..................................................................145
Margalith Galun
VIII.C. Lichenization................................................................................................................153
Margalith Galun
INDEX..........................................................................................................................................173
TABLE OF CONTENTS
Volume III
SECTION IX: CHEMICAL CONSTITUENTS OF LICHENS
IX.A. Secondary Metabolic Products..........................................................................................3
Margalith Galun and Adiva Shomer-Ilan
IX.B. Storage Products of Lichens..............................................................................................9
Philip A. J. Gorin, Madalena Baron, and Marcello Iacomini
IX.C. Pigments
IX.C.l. Carotenoids.....................................................................................................................25
Bazyli Czeczuga
IX. C.2. Phycobiliproteins.......................................................................................................35
Bazyli Czeczuga
SECTION X: PRINCIPLES OF CLASSIFICATION AND MAIN TAXONOMIC GROUPS
X. Principles of Classification and Main Taxonomic Groups..............................................41
Josef Hafellner
SECTION XI: INTERACTION OF LICHENS AND POLLUTANTS
XI. Interaction of Lichens and Pollutants.................................................................................55
Margalith Galun and Reuven Ronen
SECTION XII: OTHER USES OF LICHENS
XII. A. The Use of Lichens in Dating.....................................................................................75
John L. Innes
XII.B. Medicinal and Other Economic Aspects of Lichens.................................................93
David H. S. Richardson
XII. C. Lichens and Pedogenesis..........................................................................................109
David Jones
SECTION XIII: METHODS FOR CULTIVATING LICHENS AND ISOLATED BIONTS
XIII. Methods for Cultivating Lichens and Isolated Bionts.................................................127
Paul Bubrick
INDEX..........................................................................................................................................141
Section IX: Chemical Constituents of Lichens
IX.A. Secondary Metabolic Products
IX. B. Storage Products of Lichens
_________________________ IX.C. Pigments
