Plant Anatomy Richard Crang† Sheila Lyons-Sobaski Robert Wise Plant Anatomy A Concept-Based Approach to the Structure of Seed Plants Richard Crang† Sheila Lyons-Sobaski Department of Plant Biology Biology Department University of Illinois at Urbana-Champaign Albion College Urbana, IL Albion, MI USA USA Robert Wise Biology Department University of Wisconsin, Oshkosh Oshkosh, WI USA †Deceased ISBN 978-3-319-77208-0 ISBN 978-3-319-77315-5 (eBook) https://doi.org/10.1007/978-3-319-77315-5 Library of Congress Control Number: 2018948823 © Springer Nature Switzerland AG 2018, corrected publication 2019 This work is subject to copyright. 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Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Cover illustration: SEM of Alium sp. leaf cross section and LM of Clematis sp. stem cross-section. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Richard was a vital contributor to this project from its infancy to its final production phase. We take some comfort that he knew the book was essentially complete prior to his passing. We greatly miss our colleague and friend, and so we dedicate this book to Richard who was a courageous and venturesome guy, a brilliant scholar, an innovative teacher, and a kind, thoughtful man. SLS and RRW Preface The science of plant anatomy extends back made a substantial effort to update the to the late seventeenth century and, by subject matter, reveal new ways in which now, spans over 300 years and encom- aspects of plant anatomy play a key role in passes hundreds of thousands of reports in a variety of related disciplines in plant the scientific literature. The early plant biology, and present the topics in an anatomy research was summarized in understandable and interesting manner to 1899 by Dr. Hans Solereder in his two vol- the student and instructor. Heavy reliance ume work entitled Systematische Anatomie was made on original light and electron der Dicotyledonen: Ein Handbuch für Lab- micrographs, and color has been used oratorien der wissenschaftlichen und ange- extensively. Literature citations were kept wandten Botanik. The 1908 English to a minimum because, in today’s elec- translation by Boodle, Fritsch, and Scott tronically searchable world, a wealth of remains as fresh, informative, and useful knowledge on any topic is a mere click or today as when it was published over two away. 100 years ago. Several important texts were published in the 1950s. The two-volume This effort was started over two decades work of Metcalfe and Chalk, Anatomy of ago when a collaboration between Prof. the Dicotyledons (1950) with second edi- Richard Crang of the University of Illinois tions in 1979 (Volume 1) and 1983 (Vol- at Urbana-Champaign and Prof. Andrey ume 2), is a thorough survey of anatomical Vassilyev of the Komarov Botanical Insti- traits and features arranged by family. tute in St. Petersburg, Russia, identified the Some of the taxonomy has been rear- need for novel approaches to the teaching ranged, but the anatomical references of plant anatomy. This led to the develop- remain accurate and valuable. The year ment that used modern educational tech- 1953 saw the publication of the first edi- nologies in a searchable, compact disk tion of the classic Plant Anatomy by Kath- format that presented a traditional view erine Esau. Encyclopedic in its coverage, regarding the anatomy of temperate seed insightful in interpretation, and complete plants, their place in evolution, and taxo- in its synthesis, “Esau” (as it has been nomic relations, with a novel approach in referred to by several generations of bota- subject delivery. Although Prof. Vassilyev nists) remains a go-to reference to this day. died in 2012, his significant contributions A second edition was released in 1977 and to botany must not be overlooked. Edu- Dr. Ray Evert authored the third, revised cated in dendrology, he devoted his life to edition, published in 2006. Additionally, plant anatomy, specializing in plant secre- the 1988 Plant Anatomy by Dr. James tory structures. Dr. Vassilyev worked at the Mauseth and Dr. Avraham Fahn’s 1967 Komarov Botanical Institute and Garden Plant Anatomy (4th edition in 1990) in St. Petersburg (formerly Leningrad), belong on every plant anatomist’s book Russia, and rose to the position of Lead shelf as valuable references. Scientist at that institute. His contributions in the field of plant anatomy, and to the In 2018, plant anatomy continues to play beginnings of this project, must be noted. key roles in studies of molecular plant biology, forestry, plant pathology, plant It has been felt for some time that a new physiology, horticulture, agronomy, and a and more extensive approach to the teach- host of related botanical disciplines. There- ing of plant anatomy should be developed. fore, the authors of this plant anatomy Such plans began in 2013 and grew to resource – printed book and e-book – have include two established plant biologists VII Preface with extensive backgrounds in plant anat- ground in microscopy applications as well omy. Prof. Robert Wise of the University of as years of experience in teaching courses Wisconsin at Oshkosh and Dr. Sheila and research in plant anatomy to this Lyons-Sobaski from Albion College in effort. Michigan each bring new ideas and ­experiences to this effort in publishing. May the concepts of plant structure and Prof. Wise integrates anatomy and elec- development help open our minds to a tron microscopy with a full background in better understanding of the interrelation- plant physiology, and Dr. Lyons-Sobaski ship of life in its various forms throughout has added strength in ecology and evolu- the Earth and, perhaps, beyond. And may tion with relevance to plant anatomy. Prof. this text help, in a limited way, to aid in Emeritus Crang conveyed a lengthy back- that fuller understanding. Richard Crang† Urbana, IL, USA Sheila Lyons-Sobaski Albion, MI, USA Robert Wise Oshkosh, WI, USA †Deceased The original version of this book was revised. The correction to this book can be found at https://doi.org/10.1007/978-3-319-77315-5_20 Acknowledgments Support from the following persons has Dr. Brett Jestrow, Fairchild Tropical been invaluable to the success of this Botanic Garden, Miami ­project. With their help, the authors gained access to literally thousands of prepared Mr. Tom Perzentka, University of Wiscon- slides and specimens and were able to con- sin, Oshkosh, WI struct a library of over 10,000 images that formed the basis of this book project. Their Dr. Brian Piasecki, Lawrence University, assistance is greatly appreciated. We par- Appleton, WI ticularly wish to acknowledge and thank Dr. George Rogers, Palm Beach State Col- Eric Stannard, Senior Botany Editor, lege, Palm Beach Gardens, FL Springer for all his help and support in this project. Ms. JoAnn Stamm, Lawrence University, Appleton, WI Dr. Ranessa Cooper, Western Illinois Uni- versity, Macomb, Illinois Dr. Qiang Sun, University of Wisconsin, Stevens Point, WI Dr. John D. Curtis, University of Wiscon- sin, Stevens Point, WI This work was supported by grants from the Hewlett-Mellon Fund for Faculty Mr. John Hardy, University of Wisconsin, Development at Albion College, Albion, Stevens Point, WI MI and the University of Wisconsin Osh- kosh Faculty Development Program, Osh- Dr. James Horn, University of Wisconsin, kosh, WI. Green Bay, WI Dr. Harry Horner, Iowa State University, Ames, Iowa IX Contents I Plants as Unique Organisms; History and Tools of Plant Anatomy 1 The Nature of Plants �������������������������������������������������������������������������������������������������������������������� 3 1.1 Plants Have Multiple Pigments with Multiple Functions ���������������������������������������������������� 5 1.2 Plants Use Water, and the Properties of Water, in Unique Ways ��������������������������������������� 6 1.3 Plants Use Anabolic Metabolism to Manufacture Every Molecule Needed for Growth and Produce Virtually No Waste ������������������������������������������������������������ 8 1.4 Cell Walls Are Nonliving Matrices Outside the Plant Cell Membrane that House and/or Perform a Variety of Functions ������������������������������������������������������������������������� 9 1.5 The Plant Life Cycle Alternates Between a Haploid Gametophyte Stage and a ­Diploid Sporophyte Stage