Introducing Biological Rhythms INTRODUCING BIOLOGICAL RHYTHMS
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Introducing Biological Rhythms INTRODUCING BIOLOGICAL RHYTHMS A Primer on the Temporal Organization of Life, with Implications for Health, Society, Reproduction and the Natural Environment Willard L. Koukkari and Robert B. Sothern College of Biological Sciences University of Minnesota St. Paul, MN, USA Willard L. Koukkari Robert B. Sothern Plant Biology Plant Biology & Laboratory Medicine University of Minnesota and Pathology 250 Bio Sciences Building University of Minnesota 1445 Gortner Avenue 250 Bio Sciences Building St. Paul, MN 55108 1445 Gortner Avenue USA St. Paul, MN 55108 [email protected] USA [email protected] Library of Congress Control Number: 2005926926 ISBN-10: 1-4020-3691-4 ISBN-13: 978-1-4020-3691-0 Printed on acid-free paper. © 2006 Springer ScienceϩBusiness Media, Inc. All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer ScienceϩBusiness Media, Inc., 233 Spring Street, New York, NY 10013, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use in this publication of trade names, trademarks, service marks, and similar terms, even if the are not identified as such, is not to be taken as an expression of opinion as to whether or not they are subject to proprietary rights. Printed in the United States of America. (SPI/SBA) 987654321 springer.com “Be ruled by time, the wisest counselor of all.” —Plutarch (~46–120 AD) Greek essayist and biographer DEDICATION To my family, colleagues and friends and all who endured the many inquiries and requests and otherwise offered their patience, understanding, advice and support during the creation of this text (RBS). Preface Reading this book will not necessarily make you a better student of life. It will, however, help you to see, to hear, and to observe “that” which would otherwise escape your attention. In the present context, “that” represents the rhythms of life, which occur within us and around us. This book has been written for students and others, including teachers in the life sciences, clinical personnel, biomedical researchers, and the general public, who are interested in the topic of biological rhythms. It is not a comprehensive treatise, but a “primer” that serves to introduce individuals to the important and fascinat- ing topic regarding the rhythmic changes that affect all living organisms. During the more than three decades that we have been engaged in research and teaching the subject of biological rhythms, our students have depleted the available supply of different books1 that provided supplemental reading material for these courses. Discussions with students, colleagues, and others often led to the question “Where can I find out more about this topic?” Faced with such a situation, we embarked upon the journey of writing this book, first putting fingers to the keyboard in the fall of 1998. Since biological rhythms are a fundamental property of all life and encompass a wide range of frequencies, from seconds to a century or more, we chose to write a book that is broad in scope, covers all major groups of organisms, and discusses rhythms with periods that span the ultradian, circadian, and infra- dian domains. In many educational institutions, a biological rhythms course serves as an elec- tive for satisfying a science requirement, as a seminar or for continuing education credit. Even for those who are outside the academic disciplines, the book serves as a primer that introduces the major characteristics of biological rhythms, cites scientific results and references, and discusses the implications and applications of rhythms. Essays provide in-depth historic and other background information that relate to the main topics. 1 E. Bünning (1958) The Physiological Clock. Berlin: Springer-Verlag. Moore-Ede MC, Sulzman FM, Fuller CA. (1982). The Clocks That Time Us. Physiology of the Circadian Timing System. Cambridge: Harvard University Press. vii viii Preface We trust that the following pages will serve as an easy to understand book that will enlighten and involve the novice, while being sufficiently technical and detailed for the experienced professional. The book includes a unique chapter on self-monitoring of one’s own rhythms (autorhythmometry), which has been one of the more fascinating topics for our students in helping them to develop a greater understanding of the characteristics and role of biological rhythms in the tempo- ral organization of their own lives. Rather than focus on a few specialized areas, our aim has been to provide a book that broadly covers the field of biological rhythms, without the pretext of being completely comprehensive. For this reason, we may have overlooked cer- tain topics or missed citing some key references. If we did not delve deeply enough into an area, we trust that the material presented will sufficiently intrigue the reader to seek out additional information. New information on molecular clocks, genes, and pathways continues to accu- mulate at a rapid pace, as do findings from chronotherapy (timing of medications), investigations into sleep and its disorders, the clinical relevance of melatonin, and the effects of excess light or unusual lighting schedules on the health and survival of organisms, including humans. Names of authors and key words in the refer- ences cited provide a starting point for the search for additional information or clarification on topics of further interest. Finally, while we have tried to be as conscientious as possible in citing and writing about topics and facts contained herein, an error now and then might have occurred. In spite of the eyes of many, any such errors that slipped by our atten- tion are entirely our responsibility. We certainly welcome comments from the readers on any aspect of this book. June 2005 Willard L. Koukkari Robert B. Sothern Acknowledgments We wish to thank both old and new associates with whom we have collaborated and learned from over several decades. In addition, we are indebted to more than a hundred students who read and commented upon the many early drafts of vari- ous chapters as part of their biological rhythms coursework or independent study. We especially thank Van D. Gooch, Mary Hoff, and Tom Soulen for providing invaluable comments and suggestions to early versions of this book. We thank Kathleen Ball for checking facts in the final version against the references. We gratefully acknowledge our many colleagues, co-workers, faculty, staff, students, and friends who provided invaluable assistance with several aspects of this book, including comments, data, drawings, photos, references, and reviews. These include (in alphabetical order): Martina Abramovich, Mary Albury-Noyse, Kathy Allen, Tracy Anderson, Leonard Banaszak, Frank Barnwell, Evelyn Borchert, Robert Brambl, Ris Char- vat, Anita Cholewa, Philip Clausen, Germaine Córnelissen-Guillaume, Jayna Ditty, Jay Dunlap, Ann Dunnigan, Mark Ebbers, Linda Eels, Roger Eggen, John Erwin, Jim Fuchs, David Gartner, Steve Gantt, Florence Gleason, Stuart Gold- stein, William Gray, Francis Guillaume, Franz Halberg, David Hansen, Leon Hardman, Geoffrey Harms, Lynn Hartweck, Odette Holter, David Houghton, Roberta Humphries, Anders Johansson, Eugene Kanabrocki, Daniel Kaplan, John Kettinger, Marcia Koukkari, Mark Koukkari, Kristin Kramer, Mitch Lazar, Don Luce, Scott Madill, Julie Martinez, Krishona Martinson, C. Robertson McClung, David McGlaughlin, Robert McKinnell, Bernard Millet, Cathy Moore-Arcand, Kris Morgan, Mary Mortenson, David Mottet, Dwight Nelson, Sally Noll, Karen Oberhauser, Adeola Okusami, Neil Olszewski, Sandra Orcutt, James Orf, Dana Peterson, Richard Philips, Brian Piasecki, Nora Plesofsky, Korise Rasmusson, Anke Reinders, Tony Sanderfoot, Mark Sanders, Bill Schmidt, Janet Schottle, Michael Simmons, Carolyn Silflow, Gail Slover-Arcari, Pete Snustad, Jo Strand, Régine Terracol, John Tester, James Thompson, Edgar Wagner, John Ward, Chad Walstrom, Thomas Waters, George Weiblen, Andi Weydahl, and Michael Young. To all of those listed above go our profound and heartfelt thanks for their unstinted and cheerfully rendered help! ix Contents Preface vii Acknowledgments ix 1 The Study of Biological Rhythms Introduction ................................................................................................... 1 A Time for Everything .................................................................................. 4 Three Rhythm Domains ................................................................................ 6 Implications of Body Clocks ....................................................................... 11 Essay 1.1: Accidents and Catastrophes ................................................... 11 Chronobiology: An Integrating Discipline .................................................. 12 Chapters in This Book ................................................................................. 14 Take-Home Message ................................................................................... 15 2 General Features of Rhythms: Terminology and Characteristics Introduction ................................................................................................. 19 Details of a Rhythm..................................................................................... 20 Period