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ORCHID BIOLOGY: REVIEWS AND PERSPECTIVES, VIII
ORCHID BIOLOGY: REVIEWS AND PERSPECTIVES, VIII
Dedicated by the editors to James B. Comber, Choy Sin Hew
and Ruyso Tanaka
Orchid Biology:
Reviews and Perspectives, VIII
Edited by
TIIUKULL Department of Botany, Institute of Zoology and Botany, Agricultural University, Estonia
and
JOSEPH ARDITTI Department of Developmental and Cell Biology, University of California, Irvine
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
A C.I.P. Catalogue record for this book is available from the Library of Congress.
ISBN 978-90-481-6002-0 ISBN 978-94-017-2500-2 (eBook) DOI 10.1007/978-94-017-2500-2
Printed on acid-free paper
All Rights Reserved © 2002 Springer Science+Business Media Dordrecht OriginaIly published by Kluwer Academic Publishers in 2002 Softcover reprint ofthe hardcover lst edition 2002 No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work.
Contents
Associate editors ................................................................................ vi Board of editors .................................................................................. vi Contributors ....................................................................................... vii Persons to whom this volume is dedicated ........................................ xi Preface ................................................................................................ xv Note .................................................................................................... xv General preface ............................................................................... xvii HISTORY-BIOGRAPHY ..................................................................... 1
1 Plinius Germanicus and Plinius Indicus: Sixteenth and Seventeenth Century Descriptions and Illustrations of Orchid "Trash Baskets," Resupination, Seeds, Floral Segments and Flower Senescence in the European Botanical Literature
Ursula Wehner, Wolfgang Zierau, and Joseph Arditti MORPHOLOGY ................................................................................ 83
2 Ontogeny of Orchid Flowers Hubert Kurzweil and Alexander Kocyan
ECOLOGY ....................................................................................... 139 3 Population Dynamics of North Temperate Orchids
Tiiu Kull DEVELOPMENT ............................................................................ 167
4 Development of Orchid Seeds and Seedlings Tatyana Vinogradova and Elena V. Andronova
GENERA .......................................................................................... 235 5 Temperate Oriental Cymbidium Species
Kee-Yoeup Paek and Hosakatte Nianjana Murthy EMBRYOLOGY-SEEDS ................................................................. 287
6 Orchid Embryos Tim Wing Yam, Edward C. Yeung, Xiu-Lin Ye, Sze-Yong Zee, and
Joseph Arditti HISTORY-SEEDS ............................................................................ 387
7 Orchid Seeds and their Germination: An Historical Account Tim Wing Yam, Helen Nair, Choy Sin Hew, and Joseph Arditti
APPENDIX ...................................................................................... 505 Orchid Viruses - A Compendium
SekMan Wong Index of Persons ............................................................................... 547 Index of Organisms .......................................................................... 561 Index of Subjects .............................................................................. 574 Contents of previous volumes .......................................................... 581
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Associate Editors
Randolph S. Currah, Department of Biological Sciences, University of Alberta, Canada
Choy Sin Hew, Department of Botany, National University of Singapore, Singapore
Syoichi Ichihashi, Department of Life Science, Aichi University of Education, Japan
Gustavo A. Romero, Oakes Ames Orchid Herbarium, Harvard University, USA
Tim Wing Yam, Singapore Botanic Gardens, Singapore
Edward C. Yeung, Department of Biological Sciences, University of Calgary, Canada
Board of Editors
In addition to the editors and associate editors the following individuals took part in the preparation of this volume by providing advice, comments and reviews:
E. M. 'Monty' Beekman, University of Massachusetts, Amherst, Massachusetts, U. S. A.
Alec M. Pridgeon, Royal Botanic Gardens, Kew, U. K.
Rudolf Schmid, Department of Integrative Biology, University of California, Berkeley, California, U. S. A.
Jelte van Andel, Laboratory of Plant Ecology, University ofGroningen, Haren, The Netherlands
Jo H. Willems, Department of Plant Ecology and Evolutionary Biology, Utrecht University, Utrecht, The Netherlands
Contributors
ELENA V. ANDRONOVA is a senior scientist at the Komarov Botanical Institute in St. Petersburg, Russia. She graduated from St.Petersburg University in 1984 and received her Ph. D. in 1988 from the Department of Embryology and Reproductive Biology at the Komarov Botanical Institute. Dr. Andronova's research interests center on the reproductive structures and biology, and morphogenesis of orchids, especially northern terrestrial species. E-mail: [email protected] with "For L. Andronova" in the subject line.
JOSEPH ARDITTI received his Ph. D. from the University of Southern California (USC) in 1965 and remained there as a lecturer until 1966 when he accepted a position as Assistant Professor of Biological Sciences at the University of California, Irvine (UCI). He remained at UCI for his entire career and retired in 2001 becoming Professor of Biology Emeritus. Dr. Arditti spent many of his summers and sabbatical leaves in Indonesia (the Bogor Botanical Gardens and Flora Sari Orchids with Mr. and Mrs Soediono), Malaysia (with Professor Helen Nair at the Botany Department, University of Malaya, Kuala Lumpur) and Singapore (the Department of Botany, National University of Singapore which he considered his second academic home with Professors P. N. Avadhani, C. S. Hew and A. N. Rao) (both now retired). E-mail:[email protected].
CHOY SIN HEW is Professor (now retired) of orchid physiology in the Department of Biological Sciences at the National University of Singapore. He received his doctorate at Queen's University in Canada and was a postdoctoral
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fellow with Prof. Martin Gibbs at Brandeis University. Prof. Hew is a member of several learned societies and serves as advisor to an orchid research center in China and orchid societies in Singapore and Malaysia. He is also consultant to an orchid farm. E-mail: [email protected].
ALEXANDER KOCYAN is a graduate student working towards his Ph. D. at the Institute of Systematic Botany, University of ZUrich, Switzerland. He is studying floral structure and development of apostasioid orchids and related Asparagales. To further his studies he has taken field trips to Borneo and Tasmania. Mr. Kocyan is also involved in protection of endangered native orchid species in Switzerland. E-mail: [email protected]
TIIU KULL received her Ph. D. in botany from the University of Tartu, Estonia in 1997 with a dissertation on population dynamics of Cypripedium calceolus. She has been associated with the Institute of Zoology and Botany during her entire career. Since 1998 she has been head of the department of botany there. In addition to orchid population biology her research interests include reproductive biology of vascular plants and the protection of biodiversity. Dr. Kull has also participated in the compilation of the Flora of the Baltic Countries, Red Data books for he Baltic region and the Key-Book of Estonian plants. She also led projects on Estonian Biodiversity Strategy and Action Plan and Atlas of Vascular Plants. For the last ten years Dr. Kull has been the president of the Estonian Orchid Protection Club. E-mail: [email protected].
HUBERT KURZWEIL is a botanist at the Compton Herbarium of the National Botanical Institute, Cape Town, South Africa. He studied floral ontogeny of various orchid groups at the University of Vienna, Austria, where he received his Ph. D. in 1985. During extensive travels in south-east Asia he was able to study tropical epiphytic orchids in the field. His studies resulted in a number of research papers on the floral morphology and ontogeny of these orchids. The results were frequently included in phylogenetic analyses which were carried out jointly with Prof. H. P. Linder, University of Cape Town. Email: [email protected].
HOSAKATTE NIRANJANA MURTHY received his Ph. D. from Kamatak University, India. He joined the same university as Lecturer in Botany in 1988. At present he is a Reader in Botany there. Dr. Murthy is interested in tissue culture and conservation of orchids. He was a visiting scientist in Prof. Paek's laboratory in Korea while coauthoring the chapter on Cymbidium. E-mail: [email protected].
HELEN NAIR received her B. Sc. Honors degree from the University of Sydney, Australia and her Ph. D. from the University of Malaya (UM) while employed as an assistant lecturer there. She was appointed Professor and Chair
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of Plant Physiology at the UM Institute of Biological Sciences and is currently the Coordinator of the Division of Biotechnology at the same institute. Her research interests include the physiology, development and molecular biology of orchids and the use of biotechnology to improve the post harvest characteristics of flowers and fruits. She consults for a commercial tissue culture laboratory at the University of Malaya. E-mail: [email protected]. [email protected].
KEE YOEUP PAEK received his Ph. D. in horticulture from Kyungpook National University in Korea in 1984 with a dissertation on the factors, which affect the proliferation of protocorm-like bodies in Cymbidium. After that he joined the Department of Horticulture, Chungbuk National University where is now Professor and Director of the Research Center for the Development of Advanced Horticultural Technology. His primary research interests are the physiology and micropropagation of flowering plants. Dr. Paek is the President of the Korean Orchid Research Society and has served as a consultant to major orchid growers in Korea. E-mail: [email protected].
TATYANA N. VINOGRADOVA graduated from the Biology Department, Moscow State University in 1991 and received her Ph. D. in botany in 1999. Since 1991 she has worked at the Tropical Plants Department in the Main Botanical Garden of the Russian Academy of Sciences. Her research interests center on the anatomical and morphological characteristics orchid development under natural conditions. Since 1984 she has carried out multi-year observations on population dynamics of northern orchids at the White Sea Biological Station of Moscow University which is located at a Polar Circle latitude. E-mail: [email protected], [email protected].
URSULA WEHNER studied Latin and history at the universities of Wiirzburg and Freiburg graduating from the latter. After that she taught these subjects in a traditional German gymnasium (high school) for many years. She is now retired. E-mail: [email protected].
SEK MAN WONG received his Ph. D. in plant pathology from Cornell University. He is currently an Associate Professor in the Department of Biological Sciences at the National University of Singapore (NUS). His research interests are in the development of new methods for virus detection and the use of viral vectors for the insertion of desirable genes into plants. Prof. Wong and his team were the first to determine the nucleotide sequence of cymbidium mosaic virus and to produce a biologically active cDNA clone of it. His team was awarded the Nagoya International Orchid Congress Encouragement Prize in 1992 for their pioneering work on the use of polymerase chain reaction in the detection of orchid viruses. In 1999 he was awarded the NUS Outstanding
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University Research award. This was followed by the Meritorious Teaching Award in 2000. He is currently Sub-Dean for research at he NUS Faculty of Science. E-mail: [email protected].
TIM WING YAM graduated with a B. Sc. in Biology from the University of Southampton in England. While there he became interested in plant genetics and orchids. Dr. Yam received a M. Phil. degree in plant breeding and genetics from Cambridge University before returning to Hong Kong where he earned his Ph. D with a dissertation on the conservation of native orchids. From 1988 to 1991, he carried out postdoctoral research at the University of California, Irvine with Prof. Joseph Arditti. Since 1991 he has been a Senior Research Officer at the Singapore Botanic Gardens, in charge of the orchid breeding program for lowland tropics which involves vandaceous orchids, Dendrobium and other hybrids. This program involves vandaceous orchids, Dendrobium and other hybrids. He also administers a project designed to conserve the native orchids of Singapore by propagating and introducing these species into natural areas of the country. E-mail: [email protected][email protected].
XIU-LIN YE graduated from the Department of Biology, Jinian University, China. Presently she is a professor at the South China Institute of Botany, Academia Sinica, Guangzhou, China. Her main research interest is in reproductive biology and breeding of hybrid orchids and rice.
EDWARD C. YEUNG is a professor at the Department of Biological Sciences, University of Calgary in Canada. He received his B. Sc. Honors degree from the University of Guelph and a Ph. D. in Biology from Yale University in 1977. His primary research interests have been reproductive biology of higher plants, especially the structural and physiological aspects of zygotic and somatic embryo development. E-mail: [email protected] .
SZE-YONG ZEE received his Ph. D. from the University of Melbourne in 1969, after which he joined the University of Hong Kong as lecturer. Presently he is Professor and Head of the Botany Department there. Dr. Zee's research focuses on plant sexual reproduction and tissue culture, and the microtubule skeleton of higher plants. He is also working on several applied aspects of orchid production in Hong Kong. E-mail: [email protected].
WOLFGANG P. ZIERAU grew up in postwar Germany, first in Berlin and later Frankfurt. He received his Dr. rer. nat. (equivalent to Ph. D.) from Maximillians University in Munich with a dissertation on experimental physics of the optical properties of matter at low temperatures. Dr. Zierau spent 1975-1976 as a post-doctoral fellow with Prof. Alex Maradudin at the Department of Physics, University of California, Irvine (UCI) working on the theoretical
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properties of solid surfaces. After that he accepted a position at the WestfaelischeWilhelms-University in Muenster, Germany where his main interest is computational physics. He also continues his collaboration with Prof. Maradudin at UCI in the field of surface science. E-mail: [email protected] .
Persons to whom this volume is dedicated
JAMES B. COMBER spent 34 years in south east Asia. Following a period as a student at the Royal Botanic Gardens, Kew where he first learned to appreciate orchids Jim obtained employment in Sabah. He remained there for twelve years. Jim moved to Java after that and worked for a Swiss chemical company. I met him for the first time near the orchid enclosure at the Bogor Botanical Gardens in Indonesia. The late Djunaidi Gandawijaja, the late Saleh Idris, a number of other members of the staff and I were trying to set up an orchid laboratory by refurbishing old equipment which included a prehistoric autoclave which was heated by burning wood under its water tank; a reciprocal shaker constructed from an old electric motor, a motorcycle drive chain, bailing wire, water pipes and a discarded fluorescent light fixture; a sterile box made of cardboard, lined with aluminum foil and fitted with plastic bags which served as long gloves; and a pre world war II (or perhaps per WWI) swing balance. Jim arrived while we were scrounging for chemicals by sorting old jars and planning sorties to local pharmacies and food stores. He offered modem chemicals, dropped off a few sample bottles immediately, promised us additional help and delivered much more later. We put the laboratory into operation several days later, managed to germinate seeds and even culture several shoot tips. This would have been impossible without Jim's help. He continued to visit us after the laboratory was in operation and never failed to provide further help.
Jim spent fifteen years in Indonesia. He moved to Thailand after that, lived seven years there, retired and returned to the U. K. At present he lives in Southampton with his wife Riam, son John and daughter Elizabeth. During most of his time in south east Asia Jim spent his free days roaming the hills, forests, countryside and road sides in search orchids which he photographed and described. These studies and photographs as well as diligent work in the Kew herbarium and elsewhere served as a basis for his Wayside Orchids of Southeast Asia (1981), Orchids of Java (1990) and Orchids of Sumatra (2001).
The first book on the orchids of Java was Die Orchideen von Java by J. J. Smith published in two parts, [text in German (1905) and illustrations (1908)] E. J. Brill in Leiden, The Netherlands. The Flora of Java by C. A. Backer and R. C. Bakhuisen van den Brink which includes the orchids was published in English 60 years later. Jim's book on the Orchid of Java (published in English by the Bentham-Moxon Trust at Kew) is the first modem treatments of the orchids ofthat large orchid-rich island. The first work on the orchids of Sumatra, Enumeration of the Orchidaceae of Sumatra and Neighboring Islands in Fedde's
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Repertorium volume 32, pages 129-386 (1933) is also by J. J. Smith. Jim's newest book on the orchids of that Indonesian island is the first modem work.
CHOY SIN HEW was born in Ipoh Malaysia on 8 November 1937. He received his B. Sc. (1960) from Nanyang University in Singapore and M. Sc. (1965 at the top of his class) and Ph. D. (1967) from Queen's University in Canada. After that he spent one year (1968-1969 as postdoctoral fellow with Prof. Martin Gibbs at Brandeis University in the U. S. A. On his return to Singapore Dr. Hew was lecturer (1969-1974) and Senior Lecturer (1975-1980) in the Biology Department at Nanyang University. When the University of Singapore and Nanyang University were combined to form the National University of Singapore (NUS), Dr. Hew became Senior Lecturer in the Botany Department of the new institution. In 1982 he was promoted to Associate Professor. Eight years later he became Professor of orchid physiology and served in this position until his retirement. At present he is a Professorial Fellow at NUS. He was made Fellow of the Linnean Society of London in 1977. The National University of Singapore recognized his teaching excellence with the Best Botany Lecturer Award in 1994 and Meritorious Teaching Awards from the Faculty of Sceince in 1999 and 2000. Further recognition came to him with Singapore National Science Award in 1997 as one of the best scientists in his country.
As a research scientist Prof. Hew published more than 100 papers in international and local peer reviewed journals as well as many articles in various orchid publications, chapters in books and other contributions. He has also lectured internationally at numerous meetings and conferences. Prof. Hew is a member of several learned societies and serves as a advisor to an orchid research center in China and orchid societies in Singapore and Malaysia. He is also consultant to a large orchid cut-flower producer.-Joseph Arditti with assistance by Sek Man Wong.
RYUSO TANAKA was born in Hiroshima, Japan on 20 August 1925. He received his B. Sc. (1950) M. Sc.(1951) and D. Sc. (1961) in botany from Hiroshima Liberal Arts and Science University (now Hiroshima University). Dr. Tanaka was a Research Associate (1951-1965), Assistant Professor (1965-1968) and Professor (1968-198 Hiroshima University. When Prof. Tanaka retired from his Professorship he was awarded an Honorary Professorship. He was President of Hiroshima University from 1989 to 1993 and held the same post at Hiroshima City University between 1993 and 2000. In addition, he also held Japanese government positions with the Excutive Committee of Academic Judges (1974-1975, 1979-1987); Executive Committee of University Establishment and School Corporation (1991-1993); Executive Committee of the Japan Academic Conferences in Genetics (1978-1981), Research (1981-1984), and Plant Sciences (1981-1988) thereby contributing significantly to
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the development of Japanese education and research. Prof. Tanaka also served for many years on the Board of Directors of the Society of Chromosome Research, and the Councils of the Botanical Society of Japan, Genetics Society of Japan, Japan Mendel Society, and many other learned societies.
Dr. Tanaka developed a research interest in orchids in 1957 after a stint as postdoctoral fellow in the laboratory of Professor Haruyuki Kamemoto at the University of Hawaii. His orchid research after that centered on chromosome phylogeny; cytogenetics and karyomorphological characteristics of resting and mitotic prophase chromosomes which can be correlated with breeding compatibilities; meiotic chromosome configurations; chromosome counts of
Dedications. A. James C. Comber. B. Choi Sin Hew. C. Ryuso Tanaka. D. Vanda Miss Joaquim, the National Flower of Singapore, an orchid to which Prof. Hew has devoted much of his time.
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native Japanese orchids; and tissue-culture of shoot primordia. He was elected a member of the orchid specialists and orchid nomenclature groups of the Royal Horticultural Society in the United Kingdom. In 1987 he chaired the Lecture Program Committee of the 12th World Orchid Conference, in Tokyo. A total of 41 graduate students, many of them orchid specialists, earned their doctorates in Prof. Tanaka's laboratory. His other research interests are with Chrysanthemum and many other wild plant groups.- Katuhiko Kondo, Hiroshima University ([email protected]).
I visited Prof. Tanaka in 1969 after students riots at the University. He told me that the rioters steered clear of his laboratory because the students were aware of his martial arts proficiency. This surprised me because he is a gentle, polite, courteous and scholarly man.- Joseph Arditti
Preface
Note
Not long after publication of Orchid Biology, Reviews and Perspectives (OB) volume VII, my co-editor, Dr. Alec M. Pridgeon informed me that the pressure of other duties, especially the editing of Genera Orchidacearum (GO) will make it impossible for him to continue as co-editor and eventually editor ofthe series. Alec is an excellent orchid scientist and editor. I was sorry to that he had to leave OB, but glad that GO will be in his able hands. The first volume of GO attests to his considerable abilities and I wish him much success in the future.
Editors of orchid publications are not the most common of species (to use a botanical analogy) and finding a replacement for Alec was not easy. However I was fortunate that Dr. Tiiu Kull agreed to become my co-editor and eventually take over the series. As is obvious from the Contributors section Dr. Kull has extensive experience as both writer and editor. My interactions with her while editing this volume have convinced me she is an excellent choice. Scientifically she brings to OB an appreciation and understanding of northern terrestrial orchids, a group, which has not received as much attention as it deserves.
Another addition to OB is Dr. Tim Wing Yam who agreed to become an associate editor. Tim, who holds a position at the Singapore Botanic Gardens, will provide expertise on seed germination, hybridization, tissue culture, species and conservation.
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When I initiated OB more than a quarter of a century ago, it was not easy to convince major orchid scientists to contribute chapters. Those who were reluctant to contribute cannot be blamed since the series was an unknown in terms of quality and future prospects. I was able to publish the first volume because a number of contributors and friends believed in me and since my associates and I wrote a good part of it. All reviews except two (one of them unsigned, both inspired by a competitor) were positive and the series was launched. Still, I had to wait for about a year after OB I was published before being able to start collecting chapters for future volumes. Most potential authors responded positively and that is why volumes II and III were published within two years of each other. My associates and I contributed to these volumes also, but we did that to cover specific topics that were within our areas of expertise or when authors failed to deliver manuscripts on time (for example chapter 4 in OB III was written for OB II but was pulled out because I had too many manuscripts; it came in handy later when a projected contribution did not materialize).
Since its establishment OB has published seven volumes, nearly 3000 pages, 49 chapters and 7 appendices by 66 authors (some of which authored or coauthored more than one contribution) from 18 countries (Australia, China, Denmark, England, France, Germany, Israel, Japan, Malaysia, Netherlands, Papua New Guinea, Russia, Scotland, Singapore, Thailand, Ukraine, U. S. A, and Venezuela). There were four publishers, Cornell University Press (volumes I-IV), Timber Press (V), John Wiley and Sons (VI) and Kluwer Academic Publishers (VII). This volume adds 7 chapters, one appendix, 12 authors and five countries. The prepublication reviewers (nearly 100) were from all over the world. I had to reject only two invited chapters. In the first instance the author refused to make reviewer-recommended revisions. The second was a chapter rejected by more than the usual number of reviewers.
My practice from the outset was to follow the orchid literature and invite authors of outstanding papers in peer-reviewed journals to contribute chapters (prepublication reviewers were selected in the same manner). After volumes II and III were published, obtaining chapters was relatively easy, but some of those I invited chose not to contribute. That is why a number of prominent and reputable orchid scientists are not represented in OB. This is not to say that I contacted every orchid scientist capable of contributing. There are too many such scientists and I inevitably missed some. However I also exercised my prerogative as sole editor to invite only scientists who in my view had well earned international scientific reputations or showed promise for thr future. This simply meant that I did not invite all those who wanted to be invited. As a result there were allegations that OB was open only to my friends, associates and me. These allegations (made only after DB acquired some prestige) were spurious at best and not even remotely based on fact or reality. To counter them I indicated in the preface to volume VI (1994) that contributed chapters for future volumes would be as welcome as solicited ones.
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No chapters were contributed after my invitation, but the unfounded allegations surfaced again following publication of volume VII. My reply to these unwarranted allegations is to again invite all those who wish to submit chapters for future volumes of OB to simply do so. All suitable chapters which pass prepublication review (this point is extremely critical because all chapters will be reviewed by the editors and at least two or three reviewers; publication will depend on their recommendations) will be published in future volumes. Potential authors must keep in mind that OB publishes only critical scientific reviews, not original research papers. Contributions must follow OB format and style. Those who wish to submit chapters should contact one of the editors for instructions to authors. I chose to discuss this matter in a separate note because it arose before Dr. Kull became co-editor.
By the time this volume is published I will be retired after 35 years at the University of California, Irvine. As Professor Emeritus I will continue my association with OB, but as time goes by Dr. Kull will become the editor.
Joseph Arditti
General Preface
With only seven chapters and one appendix per book it is not possible to thematically balance every volume of Orchid Biology, Reviews and Perspectives (OB). This was clear from the outset. Therefore, the policy of OB has always been to balance the series as whole, not individual volumes. The current volume is heavy on history (two chapters), embryology (two chapters), development (three chapters) and northern terrestrial orchids (two chapters) because some of these topics were not covered adequately in previous volumes (if the total seems to exceed seven it is because some chapters cover more than one topic). As in all previous volumes every chapter was reviewed prior to being~ccepted. All chapters were also edited for language to ensure a certain degree of stylistic uniformity.
Most first chapters in previous volumes are at least partially biographic. Many are autobiographic and written by established orchid scientists past retirement age. The first chapter in this volume is also biographic and deals with major figures in orchid history. It is also somewhat of a departure because it is not an autobiography. The major figures it deals with have been dead for nearly half a millennium ( Gesner) and 300 years (Rumphius). They made major orchid discoveries, but are not as well known, as they should be. Gesner was a physicist, physician, linguist and biologist whereas Rumphius specialized in the last three in addition to being an architect. Therefore it is fitting that their biographies were written by a physicist (Zierau), a language teacher (Wehner) and an orchid biologist (Arditti).
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Flowers are the main reason for the popularity of orchids. Their ontogeny if interesting is not as well known, as it should be. Few if any have studied it in as much detail and as well and Dr. H. Kurzweil. The chapter by him and Kocyan summarized what is known about the subject.
Population dynamics is a new topic for OB. Plant population ecology has quickly developed as a science and understanding of natural processes in distribution and abundance of orchids certainly requires further attention. Dr. Kull, the new co-editor who has studied the subject for a long time, has a chapter about it. Dr. Kull has devoted much of her time to Cypripedium calceolus, which is found in both the U. S. A. and Estonia. The distribution ofthis species can be viewed as symbolizing the fact that the current co-editors of OB are from the two countries it inhabits.
For just as long most of the work by Russian scientists was unavailable in the west because of language and politics. Even when language was not a problem (the late V. A. Poddubnaya-Arnoldi had good command of English) politics during the existence ofthe U. S. S. R. prevented or at least discouraged Russian scientists from publishing in the western literature. With the demise of the U. S. S. R., publication in the west became possible and OB has taken full advantage of this. The chapter by Dr. Tanya Vinogradova and Dr. Lena Andronova is the fourth contribution by scientists from the former Soviet Union. It summarizes Russian work on embryo and seedling development which was previously unknown in the west. Orchid embryology and seedling development have been studied in Russia for a long time.
Chinese and Japanese Cymbidium species may well be the first orchids known to be cultivated extensively by humans. They were probably grown in China at least 2000 years ago. Known as Oriental Cymbidium species they are still grown in China, Japan and Korea as well as elsewhere. They have also been studied by scientists who have learned much about their physiology, seed germination, tissue culture and taxonomy. Dr. Kee-Yoeup Paek and Dr. Hosakatte Niranjana Murthy summarize what is known about them in their chapter.
Orchid embryos are unique in terms of size, structure and requirements for germination. They have been studied extensively, but information about them is scattered in time, journals and languages. Dr. Tim Wing Yam (Singapore), Dr. Edward C. Yeung (Canada), Dr. Xiu-Lin Ye (China), Dr. Sze-Yong Zee (Hong Kong), and Dr. Joseph Arditti (USA), each contributing a different perspective, bring together what is known about them.
Research on orchid seeds and their germination has a colorful and sometimes contentious history, which spans several centuries and a number of countries. Dr. Tim Wing Yam, Dr. Helen Nair, Dr. Choy Sin Hew, and Dr. Joseph Arditti who have collaborated in several areas of orchid research over the years pooled their resources to place events into perspective. OB chapters are intended to stand on their own because readers of reference volumes often pick and choose
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what they read. That is why there are repetitions in chapters one, six and seven. We are aware of the duplications and decided not to remove them.
The appendices in DB are intended to provide information for both scientists and growers. Orchid viruses have long been the bane of growers' existence and a challenge for scientists. Dr. Sek Man Wong has done a masterful job of combining both scientific and practical information in the appendix.
This volume of DB is dedicated to three leading orchid experts. They are listed in alphabetical order. A former student (Katuhiko Kondo for Professor Ryuso Tanaka) and one of the editors (for James Comber and Choy Sin Hew) wrote about them.
J. A. thanks Jean Miller, Inter Library Loan Desk, Science Library, University of California, Irvine who very competently and cheerfully located and obtained many old, rare and obscure sources and Kathryn Kjaer of the Science Library for facilitating current literature searches.
Tiiu Kull Joseph Arditti Tartu, Estonia and Irvine, California February and March 2001
