Table Of ContentBiology of Invertebrate and
Lower Vertebrate Collagens
NATO ASI Series
Advanced Science Institutes Series
A series presenting the results of activities sponsored by the NA TO Science Committee,
which aims at the dissemination of advanced scientific and technological knowledge,
with a view to strengthening links between scientific communities.
The series is published by an international board of publishers in conjunction with the
NATO Scientific Affairs Division
A Life Sciences Plenum Publishing Corporation
B Physics New York and London
C Mathematical D. Reidel Publishing Company
and Physical Sciences Dordrecht, Boston, and Lancaster
o
Behavioral and Social Sciences Martinus Nijhoff Publishers
E Engineering and The Hague, Boston, and Lancaster
Materials Sciences
F Computer and Systems Sciences Springer-Verlag
G Ecological Sciences Berlin, Heidelberg, New York, and Tokyo
Recent Volumes in this Series
Volume 89-Sensory Perception and Transduction in Aneural Organisms
edited by Giuliano Colombetti, Francesco Lenci, and Pill-Soon Song
Volume 90-Liver, Nutrition, and Bile Acids
edited by G. Galli and E. Bosisio
Volume 91-Recent Advances in Biological Membrane Studies: Structure
and Biogenesis, Oxidation and Energetics
edited by Lester Packer
Volume 92-Evolutionary Relationships among Rodents: A
Multidisciplinary Analysis
edited by W. Patrick Luckett and Jean-Louis Hartenberger
Volume 93-Biology of Invertebrate and Lower Vertebrate Collagens
edited by A. Bairati and R. Garrone
Volume 94-Cell Transformation
edited by J. Celis and A. Graessmann
Series A: Life Sciences
Biology of Invertebrate and
Lower Vertebrate Collagens
Edited by
A. Bairati
University of Milan
Milan, Italy
and
R. Garrone
Claude Bernard University
Lyon, France
Plenum Press
New York and London
Published in cooperation with NATO Scientific Affairs Division
Proceedings of a NATO Advanced Research Workshop on
the Biology of Invertebrate and Lower Vertebrate Collagens,
held June 24-28, 1984,
at the Alessandro Volta Center for Scientific Culture,
Villa Olmo, Como, Italy
Library of Congress Cataloging in Publication Data
NATO Advanced Research Workshop on the Biology of Invertebrate and
Vertebrate Collagens (1984: Alessandro Volta Center for Scientific Culture)
Biology of invertebrate and lower vertebrate collagens.
(NATO ASI series. Series A, Life sciences; v. 93)
"Proceedings of a NATO Advanced Reserach Workshop on the Biology of In
vertebrate and Lower Vertebrate Collagens, held, June 24-28, 1984, at the
Alessandro Volta Center for Scientific Culture, Villa Olmo, Como, Italy"-T.p. ver
so.
Includes bibliographies and indexes.
1. Collagen-Congresses. 2. Invertebrates-Physiology-Congresses. 3.
Fishes-Physiology-Congresses. I. Bairati, A. II. Garrone, R. III. Title. IV. Series.
QP552.C6N37 1985 591.19/245 85-16876
ISBN-\3: 978-1-4684-7638-5 e-ISBN-\3: 978-1-4684-7636-1
001: 10.1007/978-1-4684-7636-1
©1985 Plenum Press, New York
A Division of Plenum Publishing Corporation
233 Spring Street, New York, N.Y. 10013
Softcover reprint of the hardcover 1s t edition 1985
All rights reserved
No part of this book 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
PREFACE
Knowledge in the field of the biology of the extracellular
matrix, and in particular of collagen, has made considerable progress
over the last ten years, especially in mammals, birds and ln man
with respect to very important applied medical aspects.
Basic knowledge in the animal kingdom overall has increased
more slowly and haphazardly. We, therefore, considered it useful
to organize a meeting specifically devoted to the study of the
invertebrate and lower vertebrate collagens.
The NATO Scientific Division financed an Advanced Research
Workshop aimed at bringing together experts qualified in collagen
biology (with morphological, biochemical and genetic specialization)
with researchers who are currently studying collagenous tissues of
invertebrates and lower vertebrates. The Medical-Biology Committee
of the CNR-Rome and the University of Milan also supplied interest
and support for the organization of this Meeting.
The format of the workshop consisted in: 1) main lectures on
the most recent aspects of collagen biology; 2) minireviews on the
current knowledge of collagenous tissues in the various invertebrate
phyla and in fish; 3) contributed papers on particular aspects of
research in specific fields; 4) workshops on the methodology of
studying collagen.
As we had intended, the Workshop gave a comprehensive overview
of acquired knowledge and of the present state of research actlvlty.
It permitted wide interdisciplinary discussion, enabling collabora
tions to be established and new research themes to be chosen.
This volume contains the text of all the contributions presented
at the Meeting, including posters.
v
To have brought together in one volume basic knowledge, recent
experimental results and new methodologies should make this book a
useful tool for comparative analysis and for knowledge of the mech
anisms which have governed the evolution of the collagens in the
animal world.
We most sincerely thank the organ~z~ng staff of the A. Volta
Scientific Centre at Villa Olmo, where the workshop was held, all
the collaborators of our laboratories for their valuable co-operation
given during the preparation of this volume and particularly Miss
M. Properzi for her much appreciated secretarial work.
A. Bairati R. Garrone
Milano, Gennaio 1985
WILLIAM THOMAS ASTBURY AND EMMANUEL FAURE-FREMIET
In his lecture, J. Gross quite rightly recalled the importance
of the work of W. T. Astbury and E. Faure-Fremiet as regards our
knowledge of fibrous proteins, and collagen in particular.'
W. Astbury and E. Faure-Fremiet must be considered the fathers
of modern scientific research on collagen for having introduced and
applied biophysical and ultrastructural methodologies to biological
materials, for having established the major subject areas of compar
ison and evolution in the animal world, and for having gathered
around them and instructed with their teachings many skillful
researchers.
Following up on a suggestion made by J. Gross, we should like
to briefly remember the lives and work of these men so that young
researchers may be able to draw inspiration from them and go on to
perpetuate their memory and emulate their spirit.
A. Bairati R. Garrone
William Thomas Astbury was born on February 25th, 1898 at Longton
(Stoke-on-Trent, England) into a family of pottery-workers. He
studied for a degree in physics at Cambridge where he attended courses
in crystallography given by Prof. Hutchinson who, in 1921, introduced
him to Prof. Sir William Bragg at University College London. With
Bragg, Astbury worked also at the Davy-Faraday laboratory of the
Royal Institution as part of a happy working community.
His pioneering nature and the attraction of new and unknown
research fields led Astbury to accept an invitation to transfer to
the University of Leeds in 1928 in the capacity of Professor of
Physics with responsability for organizing a laboratory dedicated
to research in the field of the physics of textile fibres. In those
days it must have seemed really strange for a scholar of crystallo
graphy to want to change the direction of his career towards the
study of organic materials whose characteristics were considered
to have practically nothing to do with the precise world of crystal
lography.
Astbury worked for 33 years in Leeds, obtaining important results
in spite of the fact that for a long time the resources available to
him were extremely limited (mainly grants from the Rockefeller Foun
dation), he had to construct his own apparatus and operate in a few
rooms in an old building. After the Second War the applied importance
of the results and the presence of a highly qualifed group of collab
orators prompted the transformation of the laboratory for the physics
of textile fibres into the "Department of Biomolecular Structure"
which, after Astbury's death in 1961, bec.ame the "Astbury Department
of Biophysics".
W. T. Astbury's research activity can be schematically subdivided
into several sectors: 1) Crystallography and experimental methodology
research, 2) research on textile fibres (wool) and on fibrous pro
teins in general, 3) research on collagen, 4) research on nucleic
acids, 5) ultrastructural research on various plant and animal
tissues, 6) research of a biomedical type.
Here we are drawing attention to only some aspects of his
extremely wide field of activity.
Faced with many technical problems depending on the nature of
the biological materials, Astbury worked out some completely new
methods and built new apparatus specially to obtain better diffracto
grams. As a result of this activity great biophysical research
development began to take place in the field of the molecular
structure of paracrystalline or crystallizable components in living
material. This experimental technical ability was the result of
his earlier collaboration with Sir W. Bragg in the crystallography
field. As a physicist accustomed to the order and complexity of
crystalline structures Astbury tirelessly sought the existence of
general ordered rules in living material, analysing the form and
structure of primary constituents and their functional modifications.
The results of his research carried out between 1928 and 1935
on the keratin of wool led to the definition of the secondary poly
peptide configurations a, S and crossed S.
Using biophysical investigation together with experimentation
(physical and chemical treatments) Astbury pointed out the relation
ships between various molecular configurations, their modifications
and the variable physical properties of the tissue.
ix
The extension of his studies to other proteins (myosin and
fibrinogen) led to the concept being established of the existence
of protein groups with different molecular structures and macromo
lecular configurations. He then compared the "collagen group" to
the K-E-M-F group (keratin, epidermin, myosin, fibrinogen).
Astbury illustrated the diffractographic characteristics of
collagen fibres hypothesizing a specific conformatio~ of aminoacid
chains responsible for the peculiar distance of 2.8 A between
adjacent aminoacids.
He, therefore, widely applied comparative analysis demonstrating
the common biophysical characteristics of collagenic tissues belong
ing to different animal species: mammal tendons and cartilage, fish
scales, swim-bladder walls, byssal threads of Bivalvia, the skate
egg-capsule, sea cucumber filaments and Annelida cuticle. The studies
of various animal tissues containing keratin (feathers and reptile
scales) in addition to those of mammal hair and of collagens permitted
Astbury to lay the foundations of modern comparative research: he
outlined the idea that the differentiation of animal species might
have a common denominator in the morphology of structural proteins,
particularly fibrous ones, and that evolution could also be sought
in the mutation of molecular and supramolecular configurations.
Of equal importance were his intuitive ideas following the analysis
of transformations observed in filamentous and globular proteins
SUbjected to denaturation, tannation or crystallization. Astbury
postulated the existence of hydrogen bridges between adjacent poly
peptide chains, and maintained that the configurations assumed by
various proteins depended on the chemical nature of side chains
underlining also the importance of the localization of the hydro
phobic elements.
His analysis of the transformations of the contractile proteins
of various muscles and of fibrinogen in fibrin permitted Astbury to
hypothesize the transformation of globular into fibrous proteins and
the possibility of producing paracrystalline fibrous structures
artificially.
~1oreover, Astbury guessed the importance of the application of
biophysical and ultrastructural techniques in the industrial and
biomedical applied fields. The methodologies introduced by Astbury
were widely used in the manufacture of natural and synthethic yarns.
The use of electron microscope allowed Astbury to check directly
many of his architectural schemes deduced from diffractographic and
polaroscopic analysis, also in relation to pathological modifications
of collagenous tissues such as bone, joints and teeth.
Astbury's activity which began in the field of crystallography
achieved its best results in the biological field where Astbury's
genius showed itself in the way he exploited experimental results
x
