The Science of Photobiology PDF
Preview The Science of Photobiology
THE SCIENCE OF PHOTOBIOLOGY THE SCIENCE OF PHOTOBIOLOGY Edited by Kendric C. Smith Stanford University School of Medicine Stanford, California A PLENUM/ ROSETTA EDITION Library of Congress Cataloging in Publication Data Main entry under title: "A Plenum/Rosetta edition." The Science of photobiology. Includes bibliographical references and index. 1. Photobiology. I. Smith, Kendric C. QH515.S37 574.1'9153 77-22130 ISBN-13: 978-1-4684-1715-9 e-ISBN-13: 978-1-4684-1713-5 001: 10.1007/978-1-4684-1713-5 A Plenum/Rosetta Edition Published by Plenum Publishing Corporation 227 West 17th Street, New York, N.Y. 10011 First paperback printing 1977 © 1977 Plenum Press, New York A Division of Plenum Publishing Corporation 227 West 17th Street, New York, N.Y. 10011 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, photocophing, microfilming, recording, or otherwise, without written permission from the Publisher Foreword Although there are several excellent books covering a few of the specialized areas of photobiology, at the present time there is no book that covers all areas of the science of photobiology. This book attempts to fill this void. The science of photobiology is currently divided into 14 subspecialty areas by the American Society for Photobiology. The first 14 chapters of this book deal with those subspecialty areas, each written by a leader in the field. Chapter 15, entitled "New Topics in Photobiology," highlights areas of research that may be desig nated sUbspecialties of photobiology in the future. This book has been written as a textbook to introduce the science of photobiology to advanced undergraduate and graduate students. The chapters are written to provide a broad overview of each topic. They are designed to contain the amount of information that might be presented in a one- to two-hour general lecture. The references are not meant to be exhaustive, but key refer ences are included to give students an entry into the literature. Frequently a more recent reference that reviews the literature will be cited rather than the first paper by the author making the original discovery. Whenever practical, a classroom demonstration or simple laboratory exercise has been provided to exemplify one or more major points in a chapter. The chapters are not meant to be a repository of facts for research workers in the field, but rather are concerned with demon strating the importance of each specialty area of photobiology, and documenting its relevance to current and/or future problems of man. A great deal of research has been performed upon the effects of temperature (arctic and desert), of pressure (deep-sea diving and space flights), and of gravity (space flights) on the physiology of man, but until recently very little effort has been expended on studying the effects of light on man. Yet, light is one of the most important elements of our environment. It is hoped that this book will serve as a basic text for introductory courses in photobiology, and as a vehicle for encouraging students to enter the field of photobiology. It is also expected that this book will be of interest to scientists outside of the area of photobiology and to interested laypersons, since it is now becoming more apparent, even to the general public, that light, both natural and artificial. has important consequences to man other than just as an aid to vision. Photobiology appears to have come of age as a major scientific discipline. Kendric C. Smith Septemher 1977 v Preface The reader should be aware of the major sources of literature and information relevant to the science of photobiology. For review articles, there are two major sources: PllOtophysioloRY, Vol umes 1-8 (A. C. Giese, ed.), Academic Press, New York, covering the years 1964-1973: and Photochemical and Photobi%Rica/ Reviews, Volumes Iff (K. C. Smith, ed.), Plenum Press, New York, that was begun in 1976. For research papers, the major source is the international journal Photo chemistry and PhotobioloRJ' (Pergamon Press, London). This journal was inau gurated in 1962, and is now the official organ of the American Society for Photobiology. The American Society for Photobiology was started in 1972 (1) to promote original research in photobiology, (2) to facilitate the integration of different disciplines in the study of photobiology, (3) to promote the dissemination of knowledge of photobiology, and (4) to provide information on the photobiologi cal aspects of national and international problems. Membership in the Society is open to persons who share the stated purpose of the Society and who have educational, research, or practical experience in photobiology or in an allied scienti fic field. The name of the Society was chosen to encompass both North and South America, but members from other parts of the world are also welcome. The Journal is included in the membership dues. The American Society for Photo biology holds an annual scientific meeting (usually in June), and publishes frequent Newsletters of interest to photobiologists. Further information may be obtained by writing to the Executive Secretary, American Society for Photobiol ogy, 4720 Montgomery Lane, Suite 506, Bethesda, MD 20014. The Association International de Photobiologie sponsors an international congress on photobiology every four years. The next congress will be held in France in 1980. The Editor vii Contents Chapter 1 Phototechnology and Biological Experimentation. . . . . . 1 John Jagger Chapter 2 Spectroscopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 27 Angelo A. Lamola and Nicholas J. Turro Chapter 3 Photochemistry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Nicholas J. Turro and Angelo A. Lamola Chapter 4 Photosensitization 87 John D. Spikes Chapter 5 Ultraviolet Radiation Effects On Molecules and Cells. .. 113 Kendric C. Smith Chapter 6 Environmental Photobiology ......................... 143 Howard H. Seliger Chapter 7 Photo medicine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 175 John H. Epstein Chapter 8 Chronobiology (Circadian Rhythms) . . . . . . . . . . . . . . . . . .. 209 Beatrice M. Sweeney Chapter 9 Extraretinal Photoreception .......................... 227 Michael Menaker Chapter 10 Vision............................................... 241 Edward A. Dratz Chapter 11 Photo morphogenesis ................................ 281 Walter Shropshire. Jr. Chapter 12 Photomovement ..................................... 313 William G. Hand Chapter 13 Photosynthesis...................................... 329 David C. Fork ix x Contents Chapter 14 Bioluminescence..................................... 371 John Lee Chapter 15 New Topics in Photobiology. . . . . . . . . . . . . . . . . . . . . . . . .. 397 Kendric C. Smith Index ................................................ 419 1 Phototechnology and Biological Experimentation 1.1. Introduction.................................................................... 1 1.2. Choosing the Source .......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 1.2.1. Visible Radiation ......................................................... 3 1.2.2. Near-Ultraviolet Radiation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 1.2.3. Far-Ultraviolet Radiation .................................................. 10 1.2.4. Hazards ....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10 1.3. Narrowing the Spectral Output. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11 1.3.1. Monochromatic Sources. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12 1.3.2. Filters................................................................... 12 1.3.3. Monochromators ......................................................... 14 104. Measuring the Radiation Energy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16 1.4.1. Terminology............................................................. 16 104.2. The Thermopile........ .. ........................ .. . .. . . . . . . ............. 17 1.4.3. Electronic Photocells. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18 1.4.4. Chemical Systems (Actinometers) and Biological Systems. . . . . . . . . . . . . . . . . . . . .. 18 1.4.5. The Fluence "Seen" by a Cell................. ...................... ...... 20 1.5. Doing the Experiment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21 1.5.1. Absorption by the Sample.... ..................................... ........ 21 1.5.2. Survival Curves .......................................................... 22 1.5.3. UV Recovery Phenomena ................................................. 24 1.504. Action Spectra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25 1.6. References..................................................................... 26 1.1. INTRODUCTION(1) A great number of experiments in photobiology can be done with remarkably simple and inexpensive equipment. Nevertheless, the experimenter usually finds, after completing his initial "rough" experiments, that he must concern himself more and more with details such as the relative effectiveness of different wavelengths and the exact energies required. Consequently, there are needs for both simple and complex apparatus. One usually tries to compromise by using equipment that is only as sophisticated as is necessary for sound experimentation. John Jagger • Biology Programs. The University of Texas at Dallas, Richardson. Texas 1 2 Chapter 1 The selection of equipment to be used depends, of course, on the nature of the problem. In deciding among the enormous variety of equipment available, considerable trouble and expense may be spared if one keeps in mind the following generalizations concerning photobiological (as opposed to photochemi cal or photophysical) experimentation: (1) high intensities of light are usually required, (2) narrow bandwidths (high monochromaticities) are usually not required, and (3) irradiation times shorter than a few seconds are usually not required. Thus, the extremely high monochromaticity of a laser and the extremely short (nanosecond range) pulse of a pulsed laser are usually not required in photobiological (as opposed to photochemical) work. A few conventions should be stated at the outset. Figure 1-1 illustrates the wavelengths, photon energies, and colors in the ultraviolet (UV), visible, and near-infrared regions. For purposes ofthis chapter, we consider far-UV radiation to lie in the range of 210-300 nm, and near-UV radiation to lie in the band 300- 380 nm. We shall not be concerned with the vacuum-UV or the near-infrared regions. The width of a band of radiation will sometimes be characterized as the "half-maximum bandwidth" (bandwidth at one-half the peak of intensity), usually when discussing filters or where the transmission curve has a Gaussian form, and sometimes as the "total bandwidth" (bandwidth at -1% of the peak of intensity), usually in discussing the output of a dye laser or the complete wavelength spread in the output of a monochromator. Finally, the SI (Systeme International) units for spectral energy and power will be used (Table 1-1). TABLE 1-1. International System of Units (Systeme International d'Unites; Sl)a ampere A lux Ix calorie cal meter m candela cd milligram mg coulomb C million electron volts MeV curie Ci minute min degree Celsius °C mole mol degree Kelvin K nanometer nm degree (angle) newton N electron volt eV ohm n erg erg ohm-centimeter n-cm farad F parts per million ppm gauss G percent % gram g radian rad henry H roentgen R hertz Hz second s hour h steradian sr joule J tesla T kilogram kg torr Torr lambert L volt V liter liter watt W lumen 1m weber Wb aNote that units named for persons are not capitalized except when abbreviated. All abbreviations stand for the plural as well as the singular forms.
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