Table Of ContentELSEVIER PUBLISHING COMPANY
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With 104 illustrations and 23 tables
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COMPREHENSIVE BIOCHEMISTRY
ADVISORY BOARD
SIR RUDOLPH A. PETERS, M.C., M.D., D.Sc, F.R.S.
Emeritus Professor of Biochemistry, Oxford; Department of Biochemistry, Cambridge
Chairman
C. F. CORI, M.D., D.Sc.
Professor of Biochemistry, Massachusetts General Hospital, Boston, Mass.
J. N. DAVIDSON, D.Sc, M.D., F.R.S.
Professor of Biochemistry, The University of Glasgow, Scotland
E. F. GALE, D.Sc, F.R.S.
Professor of Chemical Microbiology, University of Cambridge
A. BAIRD HASTINGS, B.Sc, PH.D., D.SC.
Director of Biochemistry Division,
Scripps Clinic and Research Foundation, La Jolla, Calif.
E. LEDERER, PH.D., D.SC.
Professor of Biochemistry, Faculty of Science, University of Paris
F. LYNEN
Max Planck Institute for Cell Chemistry, Munich
R. NICOLAYSEN, M.D.
Professor of Nutrition Research, University of Oslo
S. OCHOA, B.A., M.D., HON.LL.D., HON.D.SC
Professor of Biochemistry, New York University School of Medicine, New York, N.Y.
J.ROCHE, D.Sc
Professor of General and Comparative Biochemistry, Collège de France, Paris
KENNETH V. THIMANN
Professor of Biology, University of California, Santa Cruz, Calif.
A. W. K. TISELIUS, D.Sc, FOR.F.R.S.
Professor of Biochemistry, Institute of Biochemistry, Uppsala, Sweden
F. G. YOUNG, M.A., D.Sc, F.R.S.
Professor of Biochemistry, University of Cambridge
C O M P R E H E N S I VE
B I O C H E M I S T RY
SECTION I (VOLUMES 1-4)
PHYSICO-CHEMICAL AND ORGANIC ASPECTS
OF BIOCHEMISTRY
SECTION II (VOLUMES 5-11)
CHEMISTRY OF BIOLOGICAL COMPOUNDS
SECTION III (VOLUMES 12-16)
BIOCHEMICAL REACTION MECHANISMS
SECTION IV (VOLUMES 17-21)
METABOLISM
SECTION V (VOLUMES 22-29)
CHEMICAL BIOLOGY
HISTORY OF BIOCHEMISTRY (VOLUME 30)
GENERAL INDEX (VOLUME 31)
C O M P R E H E N S I VE
B I O C H E M I S T RY
EDITED BY
MARCEL FLORKIN
Professor of Biochemistry, University of Liège (Belgium)
AND
ELMER H. STOTZ
Professor of Biochemistry, University of Rochester, School of Medicine
and Dentistry, Rochester, N.Y. (U.S.A.)
VOLUME 27
PHOTOBIOLOGY, IONIZING RADIATIONS
ELSEVIER PUBLISHING COMPANY
AMSTERDAM • LONDON • NEW YORK
1967
CONTRIBUTORS TO THIS VOLUME
PETER ALEXANDER, D.Sc, PH.D.
Chester Beatty Research Institute, Institute of Cancer Research : Royal Cancer Hospital,
Fulham Road, London, S.W. 3 (Great Britain)
DR. J. AMESZ
Biophysical Laboratory, University of Leiden, Schelpenkade 14A, Leiden
(The Netherlands)
C. D. B. BRIDGES, PH.D.
Medical Research Council (Vision Research Unit), Institute of Ophthalmology,
Judd Street, London, W.C.I (Great Britain)
DR. L.N.M. DUYSENS
Professor of Biophysics, Biophysical Laboratory, University of Leiden, Schelpenkade 14A,
Leiden (The Netherlands)
S. B. HENDRICKS, PH.D.
Soil and Water Conservation Research Division, Mineral Nutrition Laboratory, U.S.
Department of Agriculture, Agricultural Research Service, Beltsville, Md. (U.S.A.)
FRANK H. JOHNSON, A.B., A.M., PH.D.
Professor of Biology, Department of Biology, Princeton University, Princeton,
N.J. 08540 (U.S.A.)
JOHN T.LETT, PH.D.
Chester Beatty Research Institute, Institute of Cancer Research : Royal Cancer Hospital,
Fulham Road, London, S.W. 3 (Great Britain)
JANE K. SETLOW, PH.D.
Biology Division, Oak Ridge National Laboratory, Oak Ridge, Tenn. (U.S.A.)
H. W. SIEGELMAN, PH.D.
Crops Research Division, Plant Physiology Laboratory, U.S. Department of Agriculture,
Agricultural Research Service, Beltsville, Md. (U.S.A.)
MELVIN I. SIMON, PH.D.
Assistant Professor of Biology, Department of Biology, University of California at San
Diego, La Jolla, Calif. 92038 (U.S.A.)
KENNETH V. THIMANN, B.Sc, A.M., PH.D.
Professor of Biology and Provost of Crown College, Division of Natural Sciences,
University of California, Santa Cruz, Calif. 95060 (U.S.A.)
GENERAL PREFACE
The Editors are keenly aware that the literature of Biochemistry is already
very large, in fact so widespread that it is increasingly difficult to assemble
the most pertinent material in a given area. Beyond the ordinary textbook
the subject matter of the rapidly expanding knowledge of biochemistry is
spread among innumerable journals, monographs, and series of reviews.
The Editors believe that there is a real place for an advanced treatise in
biochemistry which assembles the principal areas of the subject in a single
set of books.
It would be ideal if an individual or small group of biochemists could
produce such an advanced treatise, and within the time to keep reasonably
abreast of rapid advances, but this is at least difficult if not impossible.
Instead, the Editors with the advice of the Advisory Board, have assembled
what they consider the best possible sequence of chapters written by
competent authors; they must take the responsibility for inevitable gaps of
subject matter and duplication which may result from this procedure.
Most evident to the modern biochemist, apart from the body of knowledge
of the chemistry and metabolism of biological substances, is the extent to
which he must draw from recent concepts of physical and organic chemistry,
and in turn project into the vast field of biology. Thus in the organization
of Comprehensive Biochemistry, the middle three sections, Chemistry of
Biological Compounds, Biochemical Reaction Mechanisms, and Metabolism
may be considered classical biochemistry, while the first and last sections
provide selected material on the origins and projections of the subject.
It is hoped that sub-division of the sections into bound volumes will not
only be convenient, but will find favour among students concerned with
specialized areas, and will permit easier future revisions of the individual
volumes. Toward the latter end particularly, the Editors will welcome all
comments in their effort to produce a useful and efficient source of biochem
ical knowledge.
M. FLORKIN
Liège/Rochester E. H. STOTZ
PREFACE TO SECTION V
(VOLUMES 22-29)
After Section IV {Metabolism), Section V is devoted to a number of topics
which, in an earlier stage of development, were primarily descriptive and
included in the field of Biology, but which have been rapidly brought to
study at the molecular level. "Comprehensive Biochemistry", with its chemical
approach to the understanding of the phenomena of life, started with a first
section devoted to certain aspects of organic and physical chemistry, aspects
considered pertinent to the interpretation of biochemical techniques and to
the chemistry of biological compounds and mechanisms. Section II has dealt
with the organic and physical chemistry of the major organic constituents of
living material, including a treatment of the important biological high
polymers, and including sections on their shape and physical properties.
Section III is devoted primarily to selected examples from modern enzymology
in which advances in reaction mechanisms have been accomplished. After
the treatment of MetaboUsm in the volumes of Section IV, "Comprehensive
Biochemistry", in Section V, projects into the vast fields of Biology and deals
with a number of aspects which have been attacked by biochemists and
biophysicists in their endeavour to bring the whole field of life to a molecular
level. Besides the chapters often grouped under the heading of molecular
biology, Section V also deals with modern aspects of bioenergetics, immuno-
chemistry, photobiology and finally reaches a consideration of the molecular
phenomena that underlie the evolution of organisms.
M. FLORKIN
Liège/Rochester E. H. STOTZ
COMPREHENSIVE BIOCHEMISTRY
Section V - Chemical Biology
Volume 22. Bioenergetics
I. Quantum biochemistry by A. PULLMAN AND B. PULLMAN. II. Mechanisms of energy
transfer by TH. FÖRSTER. III. Charge transfer in biology (a) Donor-acceptor complexes
in solution, (b) Transfer of charge in the organic solid state by F. J. BULLOCK. IV. Active
transport and ion accumulation by P. MITCHELL. Subject index.
Volume 23. Cytochemistry
I. Nucleus by G. SIEBERT. II. The nucleolus by H. BUSCH. III. Mitochondria by R. J.
BARRNETT. IV. Lysosomes by A. L. TAPPEL. V. The cell surface membrane by R. COLE-
MAN AND J. B. FINEAN. VI. Microbial cytology by M. R. J. SALTON. VII. Endoplasmic
reticulum, secretory granules and Golgi apparatus by G. PALADE. Subject index.
Volume 24. Biological information transfer. Viruses. Chemical immunology
I. Biological information transfer, (ä) DNA synthesis and replication, (b) RNA metab
olism. 1. Template RNA; 2. Transfer RNA, amino acid activation, (c) Protein syn
thesis. 1. Ribosomal function; 2. The genetic code, id) Biochemical individuality (with
general treatment of genetic errors), (e) Chemical mutagenesis. (/) Phage genetics. II.
Biochemistry of viruses. III. Immunochemistry. Subject Index.
Volume 25. Regulatory functions, membrane phenomena
I. Regulatory functions, (a) Hormone mechanisms, (b) Allosteric effects and feedback
mechanisms, (c) Enzyme induction and repression. II. Membrane phenomena, (a) Bio
electric potentials (incl. nerve impulse), (b) Secretion phenomena, (c) Cell permeability.
Subject index.
Volume 26. Extracellular and supporting structures
I. Biochemistry of the plant cell wall by S. M. SIEGEL. II. Bacterial cell walls by J.-M.
GHUYSEN, J. L. STROMINGER AND D. J. TIPPER. III. Somatic and capsular antigens of
gram-negative bacteria by O. LÜDERITZ, K. JANN AND R. WHEAT. IV. Chitinous struc
tures by CH. JEUNIAUX. V. Calcified shells by K. M. WILBUR AND K. SIMKISS. VI. Col
lagen and bone by S. FITTON-JACKSON. VII. Cartilage by A. J. BARRETT. VIII. Dental
enamel by M. J. GLIMCHER. IX. Extracellular fibrous protein: The silks by F. LUCAS
AND K. M. RUDALL. X. Intracellular fibrous proteins and the keratins by K. M. Ru-
DALL. Subject index.
COMPREHENSIVE BIOCHEMISTRY
Section V - Chemical Biology (continued)
Volume 27. Photobiology, ionizing radiations
I. Phototropism by K. V. THIMANN. II. Biochemistry of visual processes by C. D. B.
BRIDGES. III. Bioluminescence by F. H. JOHNSON. IV. Photosensitization by M. I.
SIMON. V. The effects of ultraviolet radiation and photoreactivation by J. K. SETLOW.
VI. Phytochrome and photoperiodism in plants by S. B. HENDRICKS AND H. W. SIEGEL-
MAN. VII. Photosynthesis by L. N. M. DUYSENS AND J, AMESZ. VIII. Effects of ionizing
radiations on biological macromolecules by P. ALEXANDER AND J. T. LETT. Subject
index.
Volume 28. Morphogenesis, differentiation and development
I. Fertilization by A. MONROY. II. Behaviour of nucleic acids during early development
by J. BRÄCHET. III. Biochemical pathways in embryos by E. SCARANO AND G. AUGUSTI-
Tocco. IV. Factors of embryonic induction by T. YAMADA. V. Biochemistry of am
phibian metamorphosis by R. WEBER. VI. Biochemical correlations in insect meta
morphosis by L. I. GILBERT. Subject index.
Volume 29. Comparative biochemistry, molecular evolution
I. Comparative biochemistry, (a) Basic concepts, (b) Autotrophic metabolism, (c)
Chemical needs in heterotrophs. (d) Biochemical cycles in the biosphere, (e) Biochemis
try and taxonomy. II. Molecular evolution, (a) Molecular adaptations to the physical
environment, (b) Molecular adaptations to the biological environment, (c) Heteromor-
phic aspects of molecular evolution, (d) Evolution of biochemical systems, physiologi
cal radiations, (e) Biosynthesis and phylogeny. (J) Paleobiochemistry. (g) Chemical
evolution and prebiological evolution. Subject index.
Chapter I
Phototropism
KENNETH V. THIMANN
Division of Natural Sciences, University of California,
Santa Cruz, Calif (U.S.A,)
1. Introduction
Phototropism is the orientation of the direction of growth by light. It is
almost universal in elongating green plants, and the reaction is particularly
sensitive in seedlings, which may curve towards (positive phototropism) or
away from (negative phototropism) the light source. Roots (especially of
members of the Cruciferae) sometimes show weak negative phototropism,
but because of its weakness and variability this response has scarcely been
studied. In addition to green plants a small number of fungi are phototropic,
especially the stalks of their fruiting bodies (sporangiophores).
It is important to note from the definition that phototropism is a growth
response and therefore cannot occur in a non-growing organism. However,
in a few cases, germination i.e., the beginning of growth, has its direction
determined by the direction of a light source. These phenomena are often
considered under phototropism. Free movement of motile organisms,
particularly unicells, is often oriented by light too, but because this reaction,
known as phototaxis, has few features in common with phototropism it will
not be considered here.
2. Types of phototropic response
(a) Higher plants
It is a curious fact that all of the well-studied instances of phototropism
comprise more than one reaction. The classic example of this is given by
References p. 27
[i]
