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OTHER TITLES IN THE ZOOLOGY DIVISION Vol. 1. h v e N - A n Outline of Developmental Physiology Vol. 2 . RhwN-Morphogenesis: Development The Analysis of Molluscan Vol. 3 . SAVORY-Instinctive Living Vol. 4. KEmuT-Implications of Evolution Vol. 5. TARTAR-The Biology of Stentor Vol. 6 . JENKIN-Animal Hormones-A Comparative Survey Vol. 7. ComIss-The Ciliated Protozoa Vol. 8 . GEORGE-The Brain as a Computer Vol. 9. ARTnuR-Ticks and Disease Vol. 10. RAVEN-Oogenesis Vol. 11. MA"-Leeches (Hirudinea) OTHER DIVISIONS IN THE SERIES O N PURE A N D APPLIED BIOLOGY B I 0 C H EM I S T R Y BOTANY MODERN TRENDS IN P H Y S I O L O G I C A L S C I E N C E S PLANT PHYSIOLOGY 1 W ^ ϋ^ Frontispiece Metachronal waves of membranelles of Stentor polymorphus THE BIOLOGY OF CILIA AND FLAGELLA BY M. A. SLEIGH Department of Zoology. The University, Exeter PERGAMON OXFORD · LONDON PRESS · NEW YORK 1962 · PARIS PERGAMON PRESS L T D . Headington Hill Hall, Oxford 4 & 5 Fitzroy Square, London, W.l PERGAMON PRESS INC. 122 East 55th Street, Nezv York 22, N.Y. GAUTHIER-VILLARS 55 Quai des Grands-Augustins, Paris 6 PERGAMON PRESS G.m.b.H. Kaiserstrasse 75, Frankfurt am Main Copyright © 1962 PERGAMON PRESS L T D . Library of Congress Card No. 62-19269 Set in Imprint 11 on 12 pt. and printed in Great Britain by BELL AND BAIN, L T D . . . . the general existence of the ciliary motion in the Animal Kingdom is already sufficiently established, . . . whoever has opportunities and inclination to cultivate this field of inquiry will find his labour rewarded by much curious and interesting discovery. D R . W. SHARPEY, 1835 PREFACE IMPORTANT advances in our knowledge of the structure and physiology of cilia have been made since Gray's classical monograph on Ciliary Movement was published in 1928. In particular, the improvement of techniques of electron microscopy in the last decade gave rise to a renewed interest in the structure of cilia which has been rewarded with the discovery of a widespread and consistent fibril pattern in all true cilia, flagella and sperm tails. A parallel growth of interest in the physiology of the beating and co-ordination of cilia has been taking place in these years, and has been given an added stimulus by the success of morphological studies. The primary purpose of this book is to review advances in these fields since 1928, for in these 34 years no attempt has been made to summarize fundamental work on cilia and flagella in a comprehensive way. A review by J. A. Rivera entitled Cilia, Ciliated Epithelium and Ciliary Activity was published while the present book was in the press; Rivera approached the subject in an entirely different way in summarizing the effects of various agents on ciliary activity from the point of view of the human physiologist, so that the two reviews scarcely overlap at all. In the present book modern ideas on structure are linked with modern ideas on physiology in an attempt to make a single coherent story. The field of study has been divided into four parts: ciliary structure, the effects of various agents on ciliary activity, the physiology of beating of cilia and the physiology of co-ordination. The second of these is intended for reference and for the introduction of evidence used in the third and fourth parts. Summaries of the material included will be found at the beginning of Chapter II (structure of a typical cilium) and at the ends of Chapters IV (beating activity) and V (co-ordination). The literature contains many thousands of references to ciliary structure and functioning, and, in order not to overload the book with bibliography, some selection had to b