Table Of ContentINTERNATIONAL SERIES OF MONOGRAPHS ON
PURE AND APPLIED BIOLOG Y
Division: MODERN TRENDS IN PHYSIOLOGICAL SCIENCES
GENERAL EDITORS: P. ALEXANDER and Z. M. BACQ
VOLUME 15
CILIA, CILIATED EPITHELIUM,
AND CILIARY ACTIVITY
OTHER TITLES IN THE SERIES ON PURE AND APPLIED BIOLOGY
MODERN TRENDS IN PHYSIOLOGICAL SCIENCES DIVISION
Vol. 1. FLORKIN—Unity and Diversity in Biochemistry
Vol. 2. BRACHET— The Biochemistry of Development
Vol. 3. GEREBTZOFF—Cholinesterases
Vol. 4. BROUHA—Physiology in Industry
Vol. 5. BACQ and ALEXANDER—-Fundamentals of Radiobiology
Vol. 6. FLORKIN (Ed.)—Aspects of the Origin of Life
Vol. 7. HOLLAENDER (Ed.)—Radiation Protection and Recovery
Vol. 8. KAYSER— The Physiology of Natural Hibernation
Vol. 9. FRANCON—Progress in Microscopy
Vol. 10. CHARLIER—Coronary Vasodilators
Vol. 11. GROSS—Oncogenic Viruses
Vol. 12. MERCER—Keratin and Keratinization
Vol. 13. HEATH—Organophosphorus Poisons
Vol. 14. CHANTRENNE— The Biosynthesis of Proteins
BOTANY DIVISION
Vol. 1. BOR—Grasses of Burma, Ceylon, India and Pakistan
Vol. 2. TURRILL (Ed.)—Vistas in Botany
Vol. 3. SCHULTES—Native Orchids of Trinidad and Tobago
BIOCHEMISTRY DIVISION
Vol. 1. PITT-RIVERS and TATA— The Thyroid Hormones
Vol. 2. BUSH—The Chromatography of Steroids
ZOOLOGY DIVISION
Vol. 1. RAVEN—An Outline of Developmental Physiology
Vol. 2. RAVEN—Morphogenesis: The Analysis of Molluscan Development
Vol. 3. SAVORY—Instinctive Living
Vol. 4. KERKUT—Implications of Evolution
Vol. 5. TARTAR—Biology of Stentor
Vol. 6. JENKIN—Animal Hormones
Vol. 7. CORLISS—The Ciliated Protozoa
Vol. 8. GEORGE— The Brain as a Computer
CILIA, CILIATED EPITHELIUM,
AND CILIARY ACTIVITY
i,
JOSE A. RIVERA
Naval Biological Laboratory of the School of Public Health,
University of California
at Berkeley
PERGAMON PRESS
OXFORD • LONDON • NEW YORK • PARIS
1962
PERGAMON PRESS LTD.
Headington Hill Hall, Oxford
4 & 5 Fitzroy Square, London, W.l
PERGAMON PRESS INC.
122 East 55th Street, New York 22, N Y.
1404 New York Avenue N.W., Washington 5, D.C.
PERGAMON PRESS S.A.R.L.
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PERGAMON PRESS G.m.b.H.
Kaiserstrasse 75, Frankfurt am Main
Copyright
©
1962
PERGAMON PRESS INC.
Library of Congress Card No. 61-14245
Set in Imprint 11 on 12 pt. and printed in Great Britain at
THE GARDEN CITY PRESS LIMITED, LETCHWORTH, HERTFORDSHIRE
This book is sincerely dedicated to
Dr. Carl Lamanna
Scientific Director of the Naval Biological Laboratory,
University of California,
who, by his vision and encouragement,
helped to convert this project into a reality
This book
is also sincerely dedicated to
Anne,
my wife
List of Tables
PAGE
I. Distribution of cilia 8
II. Comparison of properties of ciliary and muscular contraction 37
III. Force exerted and work done by cilia 46
IV. Influence of temperature on cilia 51
V. The effect of pure isotonic sodium salt solutions pH 7.8 upon
the epithelium of Mytilus gills 63
VI. Drugs used in the treatment of congestion of the upper
respiratory tract mucosa 73
VII. The effects of some drugs on the ciliary activity 75
VIII. The action of penicillin on ciliary motion 77
IX. Comparative ciliary activity in various concentrations of
detergents and aromatics used in dentifrices (expressed as
percentage of control speeds) 78
X. Relative toxicity for ciliated epithelium of detergents and
aromatics in absolute concentrations and in proportion to
the amounts used in dentifrices and other oral preparations 79
XL Influence of pH on depression of ciliary activity caused by
solutions of nicotine 83
List of Figures
PAGE
1. Ciliary currents on the roof of the frog's mouth 1
2. The Cilioscribe 6
3. The structure of cilia 7
4. Surface view of cells in hepatic duct of helix 10
5. Effective and recovery stroke of human nasal cilium 32
6. The nature of continuous propagation and wave propagation 33
7. Longitudinal section of cilia showing the central and the peripheral
filaments (Mag. 32,900 X) facing 36
8. Ciliary movement and streaming 39
9. Efficiency of esophageal epithelium of the frog under varying loads 44
10. The effects of temperature on the mechanical activity and on the rate of
respiration of the frontal cilia of Mytilus gills 45
11. Longitudinal section of rat tracheal epithelium (Mag. 2,220 X) facing 48
12. Longitudinal section of rat tracheal epithelium (Mag. 3,760 X) facing 49
13. The effect of temperature on the rate of movement of small particles by
frontal cilia of Mytilus 52
14. Curves showing the relation between temperature and rapidity of
movement of the esophageal cilia of frog 53
15. Air currents in the nose 57
16. The effect of hydrogen ion concentration on the mechanical activity of
cilia 62
17. Capacity of negative air ions to reverse inhibition of ciliary activity
induced by positive air ions acting on isolated rabbit trachea 68
18. Action on tracheal ciliary rate of cigarette smoke, cigarette smoke and
positive ions, cigarette smoke and negative ions 69
19. The effect on cilia of hypotonic, isotonic, and hypertonic chlorate
solutions 81
x
Preface
THIS bibliographic review was carried out with the purpose of assisting
those who would arrive at a clear understanding of the present state of
knowledge of the activity of cilia and of the role these structures play in
maintaining the normal physiology of the body, especially in the respiratory
and genital systems. The widely scattered nature of the literature in this
field, and the lack of recent reviews, are justification for an attempt to
bring together in one place all the references of work done on cilia up to the
present time.
This volume is intended to be a ready-made reference source for workers
in the fields of biological sciences and medicine. It relates the diversity of
manifestations shown by cilia and ciliated epithelium in health and disease
and the effect of the physicochemical agents studied which influence the
life and death of these structures. The survey includes an almost complete
list of references of the literature on these topics consisting of 1,335 entries
from the historical beginning of studies in the field to the time of writing.
I wish to thank the many authors to whom I am indebted for citations
from their publications. For permission to reproduce Figs. 5, 6, 8, and
IS, I am indebted to Dr. Arthur W. Proetz; for Figs. 7, 11, and 12 to
Dr. Tore Dalhamn, of Karolinska Institute; for Figs. 17 and 18 to Dr.
Albert P. Krueger, and to the Cambridge University Press, for Figs. 9, 10,
13, 14, 16, and Tables II-V.
I also wish to thank Dr. Carl Lamanna, the Scientific Director of the
Naval Biological Laboratory, University of California at Berkeley, for his
wonderful help, support, and invaluable suggestions in the preparation of
this book; Dr. Roger D. Reid, Director, Biological Sciences Division,
Office of Naval Research, for his co-operation in support of carrying on this
survey; Mrs. Mary A. Cunningham, who has done long and arduous labor
helping to prepare the bibliography; Mrs. Lois Bohannan and Mrs. Jane
Clarkin, who typed the manuscript and offered valuable suggestions, and
to Miss Doris I. Clinger, for proof reading and suggestions; in fact, to all
personnel of the Naval Biological Laboratory, University of California,
who gave of their time and help in the preparation of this volume. To all
these wonderful people, my hearty thanks and deep appreciation.
Washington, D.C. J. A. R.
xi
SECTION A: GENERAL CONSIDERATIONS
CHAPTER I
Introduction
THE term cilia refers to vibratile and nonvibratile, threadlike protoplasmic
structures which project from the free surface of certain epithelial cells.
Cilia are primordial structures, and their existence was discovered early
in the history of biology as a scientific discipline. They arise early in the
evolutionary scale of differentiated tissue and were studied long before
epithelium was discovered. Cilia are present in all groups of the animal
kingdom except the Nematodes and typical Arthropods. They are also
PARASPHENOID ^-ORBITAL CURRENT
CURRENT (STRONG) (WEAK)
FIG. 1. Ciliary currents on the roof of the frog's mouth. After Merton
[767]. From Gray [411].
present in plants. In many very primitive unicellular organisms they are
present in large numbers. So simple a protozoan as Paramecium caudatum
is equipped with 2500 cilia, and they number 10,000 in Balantidium
elongatum, the largest (approximately 70 /x by 100 /x) known protozoal
parasite of man.
In very small animals, or in the invertebrates, cilia have an important
function in contraction and locomotion. In the frog (Fig. 1), cilia and ciliary
currents help to propel small particles of food and other material into the
1
