Photobiology, the science of life and light 2nd ed l bjorn (springer, 2008)

Photobiology Second Edition Photobiology The Science of Life and Light Second Edition Edited by Lars Olof Björn Lund University Lund, Sweden Lars Olof Björn Department of Plant Physiology Lund University Sölvegatan 35 SE-223 62 Lund Sweden Lars_Olof.Bjorn@cob.lu.se Library of Congress Control Number: 2007928823 ISBN: 978-0-387-72654-0 e-ISBN: 978-0-387-72655-7 Printed on acid-free paper © 2008 Springer Science+Business Media, LLC All rights reserved This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer Science+Business Media, LLC., 233 Spring Street, New York, NY 10013, USA), except for brief excerpts in connection with reviews or scholarly analysis Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden The use in this publication of trade names, trademarks, service marks, and similar terms, even if they are not identified as such, is not to be taken as an expression of opinion as to whether or not they are subject to proprietary rights 987654321 springer.com (Drawing by Per Nilsson) Photobiology I am lying on my back beneath the tree, dozing, looking up into the canopy, thinking: what a wonder!—I can see! But in the greenery above my face, an even greater miracle is taking place: Leaves catch photons from the sun and molecules from air around Quanta and carbon atoms become bound Life, for them, has just begun The sun not only creates life, it also takes away mostly by deranging DNA Damage can be, in part, undone by enzymes using photons from the sun Summer nears its end, already ’cross the sky southward aiming birds are flying by Other birds for travel choose the night relying on the stars for guiding light Imprinted in their little heads are Gemini, Orion, Dipper, other features of the sky There is room for clocks that measure day and night, Correct for movement of the sky and tell the time for flight Deep into oceans, into caves the sun cannot directly send its waves But through intricacies of foodweb’s maze, oxygen from chloroplasts, luciferin, luciferase, at times, in place, where night and darkness seem to reign, solar quanta emerge as photons once again L.O Björn 2002 Preface I started my first photobiological research project almost exactly 50 years ago, in the spring of 1957 My scientific interest ever since has been focused on photobiology in its many aspects Because I have been employed as a botanist, my own research has dealt with the photobiology of plants, but throughout this time I have been interested in other aspects, such as vision, the photobiology of skin, and bioluminescence A first edition of the present book was published in 2002, but this second edition is much expanded and completely updated Several new authors have been recruited among my eminent colleagues It has not been possible to cover all aspects of photobiology in one volume, but I feel that we have managed to catch a fair and well-balanced cross section Many colleagues promised to help, but not all lived up to their promises To those who did, and who are coauthors to this volume, I direct my thanks; I think that they have done an excellent job Living creatures use light for two purposes: for obtaining useful energy and as information carrier In the latter case organisms use light mainly to collect information but also (e.g., by coloration and bioluminescence) for sending information, including misleading information, to other organisms of their own or other species Collection of free energy through photosynthesis and collection of information through vision or other photobiological processes may seem to be very different concepts However, on a deep level they are of the same kind They use the difference in temperature between the sun and our planet to evade equilibrium, i.e., to maintain and develop order and structure Obviously, all of photobiology cannot be condensed into a single volume My idea has been to first provide the basic knowledge that can be of use to all photobiologists, and then give some examples of special topics I have had to limit myself, and one of the interesting topics that had to be left out is the thermodynamics of processes in which light is involved Thus, this book is intended as a start, not as the final word There are several journals dealing with photobiology in general, and an even greater number dealing with special topics such as vision, photodermatology, or photosynthesis There are several photobiology societies arranging meetings and other activities And last but not least, up-to-date information can be found on the Internet The most important site, apart from the Web of Science and other scientific databases, is Photobiology Online, a site maintained jointly by the American and European Societies for Photobiology (ASP and ESP, respectively), vii viii Preface at http://169.147.169.1/POL.index.html or http://www.pol-europe.net/, where details about photobiology journals and books can be obtained The subtitle of this book may be somewhat misleading There is only one science But I wanted to point out that the various disciplines dealing with light and life have more in common than perhaps generally realized I hope that the reader will find that the same principles apply to seemingly different areas of photobiology For instance, we have transfer of excitation energy between chromophores active in photosynthesis, in photorepair of DNA, and in bioluminescence Cryptochromes, first discovered as components in light-sensing systems in plants, are involved in the human biological clock, and probably in the magnetic sense of birds and other animals, and they have evolved from proteins active in DNA photorepair The study of the photomagnetic sense of birds has, in turn, led to new discoveries about how plants react to a combination of light and magnetic fields Many colleagues have been helpful in the production of this book Two of my coauthors—Professors Helen Ghiradella and Anders Johnsson—who are also close friends, have earned special thanks, because they have helped with more chapters than those who bear their names Helen has also helped to change my Scandinavian English into the American twist of the islanders’ tounge, but we have not changed the dialect of those who are native English speakers Professor Govindjee has contributed not only with his knowledge of photobiology, but also with his great experience in editing Drs Margareta Johnsson and Helena Björn van Praagh have helped with improvements and corrections, and Professor Allan Rasmusson at our department in Lund has been very helpful when I and my computer have had disagreements I have enjoyed the friendliness and help of other colleagues in the department The staff of our biology library has been very helpful and service-minded Many others have also helped, but special thanks go to my wife and beloved photobiologist Gunvor, who has supported me during the work and put up with paper and books covering the floor in our common home; to her I dedicate those chapters of the book that bear my name Lars Olof Björn Lund, Sweden March 2007 Contents Preface vii Contributors xxi The Nature of Light and Its Interaction with Matter Lars Olof Björn 1.1 Introduction 1.2 Particle and Wave Properties of Light 1.3 Light as Particles and Light as Waves, and Some Definitions 1.4 Diffraction 1.5 Polarization 1.6 Statistics of Photon Emission and Absorption 1.7 Heat Radiation 1.8 Refraction of Light 1.9 Reflection of Light 1.10 Scattering of Light 1.11 Propagation of Light in Absorbing and Scattering Media 1.12 Spectra of Isolated Atoms 1.13 Energy Levels in Diatomic and Polyatomic Molecules 1.14 Quantum Yield of Fluorescence 1.15 Relationship Between Absorption and Emission Spectra 1.16 Molecular Geometry of the Absorption Process 1.17 Transfer of Electronic Excitation Energy Between Molecules 1.18 The Förster Mechanism for Energy Transfer 1.19 Triplet States 1.20 The Dioxygen Molecule 1.21 Singlet Oxygen Principles and Nomenclature for the Quantification of Light Lars Olof Björn 2.1 Introduction: Why This Chapter Is Necessary 2.2 The Wavelength Problem 2.3 The Problem of Direction and Shape 2.4 Biological Weighting Functions and Units 1 11 14 15 18 19 22 23 29 30 31 33 34 35 36 37 41 41 42 43 46 ix x Contents Generation and Control of Light Lars Olof Björn 3.1 Introduction 3.2 Light Sources 3.2.1 The Sun 3.2.2 Incandescent Lamps 3.2.3 Electric Discharges in Gases of Low Pressure 3.2.4 Medium- and High-Pressure Gas Discharge Lamps 3.2.5 Flashlamps 3.2.6 Light-Emitting Diodes 3.2.7 Lasers 3.3 Selection of Light 3.3.1 Filters with Light-Absorbing Substances 3.3.2 Interference Filters 3.3.3 Monochromators The Measurement of Light Lars Olof Björn 4.1 Introduction 4.2 Photothermal Devices 4.2.1 The Bolometer 4.2.2 The Thermopile 4.2.3 Thermopneumatic Devices 4.3 Photoelectric Devices 4.3.1 A Device Based on the Outer Photoelectric Effect: The Photomultiplier 4.3.2 Devices Based on Semiconductors (Inner Photoelectric Effect) 4.4 Photochemical Devices: Actinometers and Dosimeters 4.5 Fluorescent Wavelength Converters (“Quantum Counters”) 4.6 Spectroradiometry 4.6.1 General 4.6.2 Input Optics 4.6.3 Example of a Spectroradiometer 4.6.4 Calibration of Spectroradiometers 4.7 Special Methods for Measurement of Very Weak Light 4.7.1 Introduction 4.7.2 Direct Current Mode 4.7.3 Chopping of Light and Use of Lock-In Amplifier 4.7.4 Measurement of Shot Noise 4.7.5 Pulse Counting 4.8 A Sensor for Catching Images: The Charge-Coupled Device 51 51 51 51 52 53 54 55 55 56 57 58 61 62 69 69 69 69 71 72 73 73 75 76 79 80 80 80 82 84 87 87 87 88 88 88 89 Light as a Tool for Biologists: Recent Developments 93 Lars Olof Björn 5.1 Introduction 93 Contents 5.2 5.3 5.4 5.5 5.6 5.7 5.8 5.9 5.10 5.11 5.12 5.13 xi Optical Tweezers and Related Techniques 93 Use of Lasers for Ablation, Desorption, Ionization, and Dissection 95 Fluorescent Labeling 96 Abbe’s Diffraction Limit to Spatial Resolution in Microscopy 97 Two-Photon Excitation Fluorescence Microscopy 99 Stimulated Emission Depletion 100 Near-Field Microscopy 101 Quantum Dots 103 Photochemical Internalization 108 Photogating of Membrane Channels 110 Photocrosslinking and Photolabeling 113 Fluorescence-Aided DNA Sequencing 115 Terrestrial Daylight 123 Lars Olof Björn 6.1 Introduction 123 6.2 Principles for the Modification of Sunlight by the Earth’s Atmosphere 123 6.3 The UV-A, Visible, and Infrared Components of Daylight in the Open Terrestrial Environment Under Clear Skies 124 6.4 Cloud Effects 127 6.5 Effects of Ground and Vegetation 127 6.6 The UV-B Daylight Spectrum and Biological Action of UV-B 128 Underwater Light Raymond C Smith and Curtis D Mobley 7.1 Introduction 7.2 Inherent Optical Properties 7.3 Apparent Optical Properties 7.4 Estimation of In-Water Radiant Energy 131 131 132 133 134 Action Spectroscopy in Biology 139 Lars Olof Björn 8.1 Introduction 139 8.2 The Oldest History: Investigation of Photosynthesis by Means of Action Spectroscopy 141 8.3 Investigation of Respiration Using Action Spectroscopy 143 8.4 The DNA That Was Forgotten 144 8.5 Plant Vision 147 8.6 Protochlorophyllide Photoreduction to Chlorophyllide a 151 670 Index Kai proteins, in cyanobacteria biological clocks, 330 Kalanchoe, circadian clock of, 324–325 Kubelka-Munk theory application of, 20–21 computer evaluation of, 21–22 description of, 19 Lambert’s law, 19 Langley calibration method, for spectroradiometry, 86–87 Lasers for ablation, desorption, ionization, and dissection, 95–96 operation of, 56 properties of, 57 LCAO See Linear combination of atomic orbitals LDP See Long day plants LEDs See Light-emitting diodes Lei flowers, phototoxicity of, 483 Lens evolution of, 248 of human eye, 226 spectral transmission of, 523–524 Lettuce hypocotyls, growth of, 149–150 Lettuce seed, germination of, 147 LH1 complex, LH2 energy transfer to, 298 LH2 complex annihilation kinetics of, 298–299 LH1 energy transfer from, 298 in light-harvesting, 293 pigments in, 295 structure of, 293–294 Lichen planus, PUVA treatment of, 581 Lichens, history of, 275–276 Light biological clock resetting by, 321–367 clocks and light in cyanobacteria, 328–331 clocks in dinoflagellate, 331–332 description of, 321–325 in Drosophila, 344–350 fungal clocks, 338–343 in humans, 358–362 in mammals, 351–358 models, 363–367 in plants, 332–338 synchronization of, 325–328 as biologist tool, 93–116 diffraction limit in microscopy, 97–99 fluorescence-aided DNA sequencing, 115–116 fluorescent labeling, 96–97 introduction to, 93 lasers for ablation, desorption, ionization, and dissection, 95–96 near-field microscopy, 101–103 optical tweezers and related techniques, 93–95 photochemical internalization, 108–110 photocrosslinking and photolabeling, 113–115 photogating of membrane channels, 110–113 quantum dots, 103–108 stimulated emission depletion microscopy, 100–101 two-photon excitation fluorescence microscopy, 99–100 circadian clock and, 438–445 genetic approaches, 441–445 physiological approaches, 439–441 diffraction of, 7–8 energy levels of diatomic and polyatomic molecules, 23–29 isolated atoms, 22–23 generation and control of, 51–66 introduction to, 51 selection, 57–66 sources for, 51–57 heat radiation of, 11–14 human tissue effect of, 578 magnetic compass dependent on, 465–474 in animals, 465–466 inclination, 466–467 localization of, 469–470 outlook of, 474 wavelengths and irradiances for, 467–469 measurement of, 69–89 charge-coupled device, 89 fluorescent wavelength converters, 79 introduction, 69 photochemical devices, 76–79 photoelectric devices, 73–76 photothermal devices, 69–73 spectroradiometry, 80–87 very weak, 87–89 nature of, 1–39 absorption and emission spectra relation, 30–31 absorption process molecular geometry, 31–33 diatomic and polyatomic molecule energy levels, 23–29 dioxygen molecule, 36–37 electronic excitation energy transfer, 33–34 Förster mechanism, 34–35 Index introduction to, isolated atom spectra, 22–23 photon emission and absorption, 9–11 quantum yield of fluorescence, 29–30 singlet oxygen, 37–39 triplet states, 35–36 particle properties of, 1–6 photochemical reactions, 197–217 cis-trans and trans-cis isomerization, 198–211 introduction, 197–198 other photosensors, 211–217 phytochrome expression and, 421 plants and, 417–418 photoreceptors for, 418–425 polarization of, 8–9 propagation in absorbing and scattering media, 19–22 quantification of, 41–49 biological weighting functions and units, 46–49 direction and shape problems, 43–46 introduction to, 41–42 wavelength problem, 42–43 reflection of, 15–17 refraction of, 14–15 scattering of, 18–19 seed habit and, 449–450 selection of, 57–66 filters with light-absorbing substances, 58–61 interference filters, 61–62 monochromators, 62–66 sources of, 51–57 electric discharges in gases of low pressure, 53–54 flashlamps, 55 incandescent lamps, 52–53 lasers, 56–57 LEDs, 55–56 medium- and high-pressure gas discharge lamps, 54–55 sun, 51 stimulated emission of, 25–27 terrestrial daylight, 123–129 cloud effects on, 127 ground and vegetation effects on, 127 introduction to, 123 sunlight modification, 123–124 UV-A, visible, and infrared components of, 124–127 UV-B spectrum and biological action, 128–129 671 as treatment in medicine, 577–586 introduction, 577–578 photochemotherapy, 581–586 phototherapy, 578–581 underwater, 131–137 apparent optical properties, 133–134 inherent optical properties, 132 introduction to, 131 in-water radiant energy estimation, 134–137 wave properties of, 1–6 demonstration for, 619 Light amplification by stimulated emission of radiation See Lasers Light-emitting diodes (LEDs) examples of, 56 as light source, 55–56 nipple arrays and, 244 Light-harvesting carotenoids in, 303 excitons in, 299–300 in photosynthetic antennas, 293–300 Light rays, 237–238 Light-response elements (LREs), in phytochromes, 437 Light-sensing evolution of, 247 melanopsin for, 202–203 phytochrome for, 164 pigments for, 197 Light-signaling components, mutations and, 435–436 Lignin, in plant evolution, 276 Linear combination of atomic orbitals (LCAO), for chromophore energy level calculations, 158 Lipids, UV-B effects on, 516–517 Liquid crystals, structural color mechanisms of, 187 Litter, UVR effect on, 520 Living mirrors in animals, 177, 234–235 eyes with, 241–242 structural color tuning and, 177–188 introduction, 177–178 multilayer stack reflection, 183–188 single thin layer reflection, 178–183 Lock-in amplifier, for very weak light measurement, 88 Locomotor activity, in Drosophila circadian system control of, 345–347 photoreceptors for, 347–350 Lomefloxacin, phototoxicity of, 485 Long day plants (LDP) 672 Index flowering in, 438–439 promoters and inhibitors, 446–447 genetic approaches to, 441–445 circadian rhythm mutants, 444–445 photoreceptors, 442–444 photoreceptors and circadian rhythm, 444 physiological approaches of, 439–442 LREs See Light-response elements Luciferin aequorin v., 600 for ATP assays, 607 in firefly bioluminescence, 599 Lumen, for luminous flux, 47 Luminous flux, 47 Lupus vulgaris, short-wavelength light for, 577–578 Lux, for illuminance, 47 Macula lutea, in spectral tuning, 167, 170 Magnetic compass, light-dependent, 465–474 inclination for, 466–467 localization of, 469–470 mechanisms of, 470–473 outlook of, 474 wavelengths and irradiances for, 467–469 Magnetic field, in light polarization, Magnetoreception chemical, 471–472 cryptochrome involvement in, 471–473 light involvement in, 465 localization of, 469–470 mechanisms of, 470–473 outlook of, 474 radical pair mechanism of, 471–472 diagnostic tools for, 473 Malate, in C4 metabolism, 272 Maleimide, for photogating of nerve cells, 111–112 Malignant melanoma (MM) description of, 563 of human skin, 560 rates of, 563–564 risk factor for, 564 Mammals biological clock in, 351–358 clocks outside SCN, 357–358 photoreceptors in retina, 353–355 pineal organ, melatonin, and photoperiodism, 355–357 SCN and pathways of, 351–353 provitamin D in skin of, 533–534 Manganese, photochemical oxidation of, 263–264 Mantis shrimp, vision of, 171 Matter, light interaction with, 1–39 absorption and emission spectra relation, 30–31 absorption process molecular geometry, 31–33 diatomic and polyatomic molecule energy levels, 23–29 diffraction of, 7–8 dioxygen molecule, 36–37 electronic excitation energy transfer, 33–34 Förster mechanism, 34–35 heat radiation, 11–14 introduction to, isolated atom spectra, 22–23 particle and wave properties of, 1–7 photon emission and absorption, 9–11 polarization of, 8–9 propagation in absorbing and scattering media, 19–22 quantum yield of fluorescence, 29–30 reflection, 15–17 refraction, 14–15 scattering, 18–19 singlet oxygen, 37–39 triplet states, 35–36 MED See Minimal erythema dose Medicine, light treatment in, 577–586 introduction, 577–578 photochemotherapy, 581–586 phototherapy, 578–581 Melanin, in skin pigmentation, 558 Melanoma, of eyes, 524–525 Melanophores, in frogs, 202 Melanopsin, in biological clock, 202–203 Melatonin activity of, 356–357 in mammalian biological clock, 355–357 for synchronizing circadian systems, 361 synthesis of, 356 Membrane depolarization, pharmacological study of, 437–438 Mercury lamps, 54 Metabolic cycles, for carbon dioxide assimilation, 270–271 C4 Metabolism atmosphere ans, 273–274 CAM v., 275 description of, 272–273 evolution of, 273 C3 Metabolism, C4 metabolism v., 272–273 Metarhodopsin, formation of, 202 Methane, oxidation of, 278 8-Methoxypsoralen (MOPS) Index medical treatment with, 484 phototoxicity of, 479–480 Methylene blue, for PDT, 584 Microdessication, by lasers, 96 Microorganisms, UVR affect on, 144–146 demonstration for, 637–638 Microscopy, diffraction limit in, 97–99 Microwave radiation, gas discharge lamp as source of, 54 Mie scattering, 18–19 Minimal erythema dose (MED), determination of, 559 Mirrors See Living mirrors MM See Malignant melanoma Models, wave and particle of light as, 5–6 Molecules electronic excitation energy transfer between, 33–34 energy levels in, 23–29 Monochromators description of, 62–63 diagram of, 65 diffraction pattern from, 63–64 gratings in, 64–66 as light filters, 62–66 slits in, 65–66 MOPS See 8-Methoxypsoralen "Multibank" retina, of birds and fish, 235 Multilayer stack, reflection in, 183–188 Nalidixic acid, phototoxicity of, 485 Nanda Hamner experiments negative responses in, 404–405 on photoperiodism and circadian rhythms, 396, 398 Natural light polarization of, 8–9 wavelength of, Near-field microscopy, 101–103 Near-field scanning optical microscopy (NSOM), 101–103 Neonatal jaundice causes of, 580–581 visible light phototherapy for, 581 Nerve cells, photogating of, 110–113 Neurospora crassa biological clock study in, 338 circadian system of, 338–341 entrainment of, 341–342 Newts magnetic compass orientation in, 467–468 magnetoreception process of light in, 465 localization of, 469–470 673 Nickel sulfate, as light filter, 60 Night-break responses (NB), in SDP and LDP, 440–442 Night interruption experiments, on photoperiodism and circadian rhythms, 397–399 Nipple arrays applications of, 244 on insect eyes, 240–241 Nitrogen monoxide, in firefly bioluminescence, 604–606 NMSC See Nonmelanoma skin cancer Nonmelanoma skin cancer (NMSC), in human skin, 561–563 Normalized frequency, 238–239 NSOM See Near-field scanning optical microscopy Nucleic acid, importance of, 145–146 Oilbirds, eye structure of, 234–235 OLEDs See Organic light-emitting diodes Ommatidia, reflective optics in, 242–243 Optical tweezers description of, 94–95 and related techniques, 93–95 Optical washboard, 94–95 Organic light-emitting diodes (OLEDs), 56 Ovarian diapause, Drosophila induction of, 408–411 Oxidative quenching, by carotenoids, 303 Oxygen deleterious effects of, 277–278 di-, 36–37 in firefly bioluminescence, 604–606 historical levels of, 278 for PDT, 585 from photosynthesis, 277 in photosynthesis, 262–265 singlet, 37–39 demonstration for, 620 Oxygenic photosynthesis, chlorophyll a in, 161 Oxygen tension, SR under, 204–205 Ozone action spectroscopy of layer of, 153 description of, 504 from photosynthesis, 277 sunlight modification by, 123–124 xenon lamp creation of, 55 Ozone layer depletion of, 506–508 human eye effects of, 523–525 introduction to, 503–504 location of, 507–508 674 Index organism influences, 520–523 process of, 506–507 description of, 504–506 PABA See Para-aminobenzoic acid PAHs See Polycyclic aromatic hydrocarbons PAR See Photosynthetically active radiation Para-aminobenzoic acid (PABA), phototoxicity of, 485–486 Particle properties, of light, 1–6 definitions, 6–7 PAS domain of phytochrome, 208 of PYPs, 205–207 PCB See Phycocyanobilin PDT See Photodynamic therapy Peas, circadian clocks in, 337 PEPC See Phosphoenolpyruvate carboxylase Perfumes, phototoxicity of, 486 PFB See Phytochromobilin Pharmacological study of aequorin, 438 of membrane depolarization, 437–438 of PHY, 437 Phase response curve for biological clock synchronization, 326–327 in humans, 360 for diapause induction overt "indicator" rhythms, 403–404 transient or non-steady-state entrainment, 401–402 Phase shifts, of circadian clocks, 365 Phenanthrene absorption spectra of, 489 phototoxicity of, 490 Pheophorbide, in dragonfish, 603 Phosphoenolpyruvate carboxylase (PEPC) in C4 metabolism, 272 in CAM, 274–275 Phosphoglycolic acid in C3 metabolism, 272 formation of, 272 Photoactive yellow proteins (PYPs) PAS domain of, 205–207 as photoreceptors, 205–207 Photoageing, of human skin, 559–560 Photoallergic contact dermatitis, 572 Photoautotrophic bacteria, near deep sea vents, 261 Photobiology of human skin, 553–572 CHS and DTH, 556–557 immunosuppression, 565–570 introduction, 553–554 photoageing, 559–560 photocarcinogenesis, 560–565 photodermatoses, 570–572 pigmentation and sunburn, 557–559 skin immune system, 555–556 skin structure, 554–555 UVR effect on, 557–558 of vitamin D, 531–546 introduction, 531–532 Photocarcinogenesis, of human skin, 560–565 animal studies of skin cancer, 564–565 malignant melanoma, 563–564 nonmelanoma skin cancer, 560–563 Photocell photon counting with, young’s double slit experiment with, 2–6 Photochemical devices, for light measurement, 76–79 Photochemical internalization advantages of, 109 description of, 108–109 methods of, 109–110 Photochemical reactions, in light perception and regulation, 197–217 cis-trans and trans-cis isomerization, 198–211 introduction, 197–198 other photosensors, 211–217 Photochemistry, in photosynthesis, 156–157 Photochemotherapy description of, 578 extracorporeal, 582–584 mechanism of action, 583–584 principle and area of use, 582–583 implementation of, 582 PDT, 584–586 PUVA, 581–582 Photochemotherapy with UV-A (PUVA), treatment with, 581–582 Photochromicity, of chromoprotein, 163 Photoconductive cells, 75 Photocrosslinking, 113–115 Photocytes, firefly control of, 604–606 Photodermatoses, of human skin, 570–572 cutaneous porphyrias, 571–572 genodermatoses, 570 idiopathic, 571 photoallergic contact dermatitis, 572 Photodiodes, 75 Photodynamic therapy (PDT) description of, 578 physiological mechanism of, 584–585 Index with porphyrins and chlorins as photosensitizers, 584–586 Photoelectric devices inner photoelectric effect, 75–76 for light measurement, 73–76 outer photoelectric effect, 73–75 Photofrin, for PDT, 585–586 Photogating advantages of, 111–113 of membrane channels, 110–113 methods of, 110–111, 113 Photographic film for light measurement, 76 young’s double slit experiment with, 2–4 Photolabeling, 113–115 Photolyases cryptochromes from, 197 for DNA repair, 513–514 evolution of, 514–515 Photomorphogenesis, 417–451 concluding remarks on, 451 demonstration of, 638–640 description of, 417–418 introduction to, 417–418 in natural environment, 447–451 energy capture improvement, 448–449 light and seed habit, 449–450 unfavorable condition avoidance, 450–451 photoreceptors for, 418–425 cryptochromes, 423–424 germination role, 426 others, 425 phototropins, 424–425 phototropism, 429 physiological roles, 425–431 phytochromes, 418–423 seedling establishment, 427–429 shade avoidance, 430–431 signal transduction, 431–438 Photomultipliers diagram of, 74 for light quantization, 42, 73–75 of very weak light, 87–89 operation of, 73–74 as photocell, ranges of, 74–75 Photon(s) blackbody radiation and, 12 emission and absorption statistics of, 9–11 energy of, irradiance and fluence rate of, 43–44 Photonic crystals examples of, 186–187 overview of, 186 675 Photooxidation of lipids, 517 of PAHs, 491 Photoperiodic induction, of circadian clocks, 366 Photoperiodic responses description of, 418 in LDP, 441–442 in SDP, 440–442 Photoperiodism circadian system with, 396–403 diapause induction, 401–403 Nanda Hamner experiments, 396, 398 night interruption experiments and Bünsow protocol, 397–399 skeleton photoperiods and bistability phenomenon, 397, 400–401 in fungi, 343 in insects and other animals, 389–411 circadian system in, 396–403 diapause and seasonal morphs, 391–392 Drosophila diapause induction, 408–411 "hourglass" alternative, 404–405 introduction, 389–390 models for, 393–396 overt "indicator" rhythms for, 403–404 photoreception and clock location, 405–408 in mammals, 355–357 models for, 393–396 in plants, 337–338, 417–451 concluding remarks on, 451 introduction, 417–418 light and circadian clock, 438–445 in natural environment, 447–451 signaling in, 445–447 Photopigments, for biological clock synchronization, 325–326 Photoreactivation of CPDs, 513–514 by UVR, 635–637 Photoreceptor(s) for biological clock synchronization, 325 in Drosophila, 347–350 in fungi, 342–343 in humans, 359–360 in mammalian retina, 353–355 in plants, 334–336 in cyanobacteria biological clocks, 328–330 definition of, 197 in dinoflagellate biological clocks, 332 mutants and circadian clocks and, 442–444 signal transduction of, 432–436 676 Index for photoperiodism, 405–408 in plants, 418–425 cryptochromes, 423–424 germination role, 426 others, 425 phototropins, 424–425 phototropism, 429 physiological roles of, 425–431 phytochromes, 418–423 seedling establishment, 427–429 shade avoidance, 430–431 signal transduction, 431–438 PYPs as, 205–207 signal transduction of, 431–438 expression profiling, 436–437 mutants and interacting factors, 432–436 pharmacological approaches, 437–438 primary reactions, 431–432 temporal differences in, 429 Photoreversibility, of phytochrome, 422–423 Photosensitizers bonding of, 482 for PDT, 584–586 for photochemical internalization, 108–109 Photosensors cryptochromes, 211–212 phototropin, 213–215 plant UV-B receptor, 215–217 Photosynthesis action spectroscopy investigation of, 141–143 afterglow of, 608–610 biological clock of, 321–322 carotenoids in, 306–307 evolution of, 255–280 ATP-synthesizing enzymes, 275 C4 metabolism, 272–274 carbon dioxide assimilation, 270–272 concluding remarks on, 280 crassulacean acid metabolism, 274–275 cyanobacteria to chloroplasts, 265–267 domains of life, 258–259 first photosynthesis, 260–261 impact on biospheric environment, 277–280 introduction to, 256 journey onto land, 275–277 oxygenic, 262–265 photosynthetic pigments and chloropast structure, 267–270 predecessors of first photosynthetic organisms, 259–260 overview of, 255, 289 photochemistry in, 156–157 review of, 257–258 spectral tuning in, 155–156 Photosynthetically active radiation (PAR) description of, underwater, 131 depth v., 135–137 Photosynthetic antennas, light-harvesting and energy transfer in, 293–300 between strongly coupled chromophores, 296–298 theoretical considerations, 294–295 between weakly dipole-coupled chromophores, 295–296 Photosynthetic charge separation, 300–303 primary electron transfer mechanism, 301–303 structure and function of bacterial reaction center, 300–301 Photosynthetic organisms, introduction to, 289–293 Photosynthetic photon flux density (PPFD), 47–48 Photosynthetic reaction center, charge separation in, 300–303 Photosynthetic unit (PSU) intercomplex excitation transfer in, 298–300 of plants, 292 of purple bacteria, 291–292 Photosystem II (PSII) antenna complexes with, 312 delayed light emission with, 608 Photosystem I (PSI), structure of, 292 Photosystems evolution of, 262–263, 268 overview of, 289–293 oxygenic photosynthesis v., 264 Phototherapy description of, 578 implementation of, 582 UV-A, 580 UV-B, 578–580 visible light, 580–581 Photothermal devices bolometer, 69–71 for light measurement, 69–73 thermopile, 71–72 thermopneumatic devices, 72–73 Phototoxicity drugs and cosmetics, 485–486 of fungal plant parasites, 484–485 introduction to, 479–481 third type, 481 type I, 479–481 type II, 480–481 Index metabolic disturbances and, 487–489 in plant defense, 482–484 of poly cyclic aromatic hydrocarbons, 489–495 to aquatic biota, 493–495 in aquatic systems, 492–493 mechanisms of, 490–491 nature and occurrence of, 489–490 Phototransistors, 75 Phototropic responses, 417–418 Phototropin(s) for biological clock synchronization, in plants, 334–336 chromophores of, 213–215 description of, 213 mutants and interacting factors of, 433–435 in phototropism, 429 in plants, 424–425 primary reactions of, 432 Phototropism, of seedlings, 429 PHY See Phytochrome Phycobiliproteins, 162–164 as antenna pigment, 162 chlorophyll and evolution of, 188 types of, 162 Phycobilisomes, 162–164 chromatic adaptation of, 164–166 in cyanobacteria, 266, 268 energy transfer in, 33–34 structure of, 290–292 Phycochrome, in cyanobacteria, 209–210 Phycocyanins absorption bands of, 164 in cyanobacteria, 164–165, 266 adjustments of, 209 Phycocyanobilin (PCB), 163 of phytochrome, 423 Phycoerythrin, in cyanobacteria, 164–165, 266 adjustments of, 209 Phycoerythrobilin, 163 Phycourobilin, 163 Phycoviolobilin, 163 Phylloerythrin, phototoxicity of, 487–488 Phytochrome (PHY) absorption spectra of, 149–150 for biological clock synchronization, 325–326 in plants, 334–336 chlorophyll and, 188 demonstration for, 640–642 discovery of, 147, 207 formation of, 147–149 light-sensing function of, 164 LREs in, 437 677 mutants and interacting factors of, 433–434 optical properties of, 208 PAS domain of, 208 pharmacological study of, 437 plant identification of, 165 in plants, 207–208, 418–423 chromophores of, 422–423 expression and localization, 421–422 gene/protein structure, 421–422 genes and gene surveys, 419–421 isolation, 418–419 seedling establishment, 427–429 primary reactions of, 431–432 spectral tuning in, 155–156 Phytochromobilin (PFB), of phytochrome, 422–423 Phytoluminography, 608–610 Pigmentation See Complexion Pigments antenna, spectral tuning in, 162 differentiation of, 289–290 light-sensing, 197 for photogating of nerve cells, 111 in photosynthesis, 142 evolution of, 267–270 in phototoxicity, 481 spectra determination of, 157–159 in spectral tuning, 155 evolutionary pressures on, 170–171 structural colors with, 187 visual description of, 166 structures of, 166–167 Pineal organ, in mammalian biological clock, 355–357 Planck’s constant, 12 Planck’s radiation law application of, 12–13 blackbody radiation for, 12–13 daylight and, 156 derivation of, 11–12 heat radiation and, 11, 31 Plane of incidence, in light reflection, 15–16 Plane of polarization, Plane-polarized light, 8–9 Plant(s) C4, 272–274 CAM in, 274–275 chlorophyll a in, 161 circadian systems of, photoperiodic time measurement, 400 color of, 156–157 demonstration for, 618–619 cryptochromes in, 211–212, 423–424 678 Index evolution of, 188, 269, 276 light acclimation of, 629–635 light effects on biological clocks in, 332–338 importance of, 333–334 mechanism and clock-controlled genes, 336–337 photoperiodism, 337–338 photoreceptors, 334–336 in natural environment, 447–451 energy capture improvement, 448–449 light and seed habit, 449–450 unfavorable condition avoidance, 450–451 photomorphogenesis in, 417–438, 447–451 concluding remarks on, 451 introduction, 417–418 in natural environment, 447–451 photoreceptor physiological roles, 425–431 photoreceptors, 418–425 photoreceptor signal transductance, 431–438 photoperiodism in, 438–451 concluding remarks on, 451 introduction, 417–418 light and circadian clock, 438–445 in natural environment, 447–451 signaling in, 445–447 photoreceptors in, 418–425 cryptochromes, 423–424 germination role, 426 others, 425 phototropins, 424–425 phototropism, 429 physiological roles of, 425–431 phytochromes, 418–423 seedling establishment, 427–429 shade avoidance, 430–431 signal transduction, 431–438 phototoxicity for defense of, 482–484 phototropins in, 424–425 phytochrome in, 207–208, 418–423 identification in, 165 provitamin and vitamin D in physiological effects of, 541 roles of, 541–542 PSU of, 292 purple bacteria v., 312–313 reaction center of, 303 rigidity of, 276 rubisco in, 259 UV-B molecular effects on, 508–510 receptors of, 215–217, 518 vitamin D in, 535, 540–541 C4 Plants, evolution of, 272–274 Plastocyanin, photosynthesis in, 258 Plastoquinone, photosynthesis in, 258 PLE See Polymorphic light eruption Plexiglas for containing light filters, 60 as light filter, 58 Poisson distribution, 10 Polarization absorption and, 32 with birefringent mediums, 15 of light, 8–9 Polyatomic molecules, energy levels in, 23–29 Polycyclic aromatic hydrocarbons (PAHs) nature and occurrence of, 489–490 phototoxicity of, 489–495 to aquatic biota, 493–495 in aquatic systems, 492–493 mechanisms of, 490–491 Polyenes complication with, 159 conjugated double bonds in, 158 excitation states of, 309 spectral tuning with, 158–159 Polymorphic light eruption (PLE), 571 Porphin, molecular structure and absorption of, 159–160 Porphyria, in humans, 487–488 Porphyrins chlorophyll from, 157 for PDT, 584–586 Porphyropsins, in dragonfish, 603 Potassium dichromate, as light filter, 58 PPFD See Photosynthetic photon flux density Previtamin D chemical reactions of, 533–534 demonstration for, 624–625 vitamin D conversion of, 534–535 Primary electron transfer, mechanism of, 301–303 Propagation mode theory, of compound eyes, 237–240 Airy disk, 239–240 geometrical optics, 238 normalized frequency, 238–239 ray optics, 238 Protein chromophore attachment to, 163–164 spectral tuning with, 167 UV-B effects on, 518 visual pigments influence of, 168–169 Protochlorophyllide photoreduction to chlorophyllide a 151–152 demonstration of, 625–627 Index Protoporphyrin IX, phototoxicity of, 487 Protozoa, bioluminescence of, 592 Provitamin D absorption spectra of, 535 chemistry and photochemistry of, 532–536 demonstration for, 624–625 1,25-dihydroxyvitamin D from, 536 in mammalian skin, 533–534 plant kingdom distribution of, 540–541 PSI See Photosystem I PSII See Photosystem II Psoriasis expression of dysfunction of differentiation, 579 hyperproliferation, 579 T-cell inflammatory reaction, 579 PUVA treatment of, 581 UV-B phototherapy for, 578–580 PSU See Photosynthetic unit Pteridines, as antenna pigment, 162 Pterin, of cryptochrome, 423–424 Pterocarpans, as type I phototoxins, 482 Pterygium, of eyes, 525 Pulse counting, for very weak light measurement, 88–89 Purine, UVR transformation of, 512 Purple bacteria photosynthesis in, 260 plants v., 312–313 PSU of, 291–292 PUVA See Photochemotherapy with UV-A PYPs See Photoactive yellow proteins Pyrene absorption spectra of, 489 phototoxicity of, 490 Quantification, of light, 41–49 biological weighting functions and units, 46–49 direction and shape problems, 43–46 introduction to, 41–42 wavelength problem, 42–43 Quantum counters See Fluorescent wavelength converters Quantum dots absorption spectrum of, 104–105 advantage of, 105 applications of, 106–107 description of, 103 emission wavelength of, 104–105 fluorescence yield of, 106 materials for, 103–104 properties of, 106 types of, 107–108 679 Radiance, 46 Radiant excitance, 46 Radiant intensity, 46 Radical pair mechanism, of megnetoreception, 471–472 cryptochromes in, 471–473 diagnostic tools for, 473 Raman scattering description of, 18–19 for lasers, 57 Rapid eye movement (REM), circadian system and, 358 Rayleigh scattering by atmosphere, 123 description of, 18–19 Reaction center bacterial excitation of, 302 structure and function of, 300–301 electron transfer in, 302–303 photosynthetic charge separation in, 300–303 primary electron transfer in, 301–303 of plants, 303 Reactive oxygen species (ROS) formation and effects of, 515–516 lipids and, 517 Red alga chromatic acclimation in, 142–143 fossils of, 266–267 phycobilisomes in, 268 Red light blue light effects v., 214 chromatic adaptation under, 165–166 far-red light antagonism with, 147–148 receptor evolution, 170, 188 Red-shifts, of chlorophyll, 161 Reflection of light, 15–17 in multilayer stack, 183–188 chemical compounds for, 185 computation of, 183–186 liquid crystals, 187 photonic crystals, 186–187 optics of, 241–242 refractive index and, 16–17, 177–178 scattering v., 18 in single thin layer, 178–183 obtaining zero reflection, 182–183 phase relations in, 179–180 reflectance calculation, 180–182 with snow cover, 127 680 Index Reflection gratings, in monochromators, 64–65 Refraction for human eye, 223–226 of light, 14–15 in homogeneous sphere, 227–228 optics of, 240–241 scattering v., 18 Refractive index description of, 14–15 of eye in water problem, 227 solution, 228–230 through a glass sphere, 227–228 of human eye, 225–226 reflection and, 16–17 variable, 228–230 wavelength v., 15 Reindeer lichen, vitamin D in, 544–545 Reproduction, bioluminescence in, 593–594 Respiration, action spectroscopy investigation of, 143–145 Retene absorption spectra of, 489 phototoxicity of, 490 11-cis-Retinal in humans, 167–168 isomerization of, 201–202 as visual pigment, 166–167 Retinal proteins, spectral tuning of, 169 Retina, mammalian circadian clock in, 357–358 SCN v., 358 Rhabdom Airy disk of, 239–240 light propagation to, 239–240 Rhodopsins demonstration of, 623–624 description of, 166, 200 in dragonfish, 603 evolution of, 248 human, 167–168 isomerization of, 200–203 energy barrier in, 199 occurrence of, 246–247 for photogating of nerve cells, 111 Rhodovibrin, energy transfer of, 310 Rickets description of, 531–532 sunlight curing of, 533 Rods of oilbirds, 234–235 in vertebrate vision, 166 of whales, 171 ROS See Reactive oxygen species Rose, cornflower color v., 172–173 Rotational energy in molecules, 23 vibrational and electronic energy v., 23 Rubisco for carbon dioxide reduction, 259 evolution of, 271–272 Rydberg constant, in atomic spectra, 22 SAD See Seasonal affective disorders Sanger method, of DNA sequencing, 115 Saponins, geeldikkop from, 488 Scalar irradiance for photobiology, 133–134 wavelength v., in water, 135–136 Scallop, refractive and reflective optics of, 241–242 Scanning eyes, of crustaceans, 242–245 Scattering in biological world, 187 effect of, 20–21 of light, 18–19 reflection and refraction v., 18 Scattering media, light propagation in, 19–22 SCC See Squamous cell carcinoma Scleroderma, UV-A phototherapy for, 580 Sclerosis, extracorporeal photochemotherapy for, 584 SCN See Suprachiasmatic nucleus SDP See Short day plants Seasonal affective disorders (SAD) human circadian rhythms and, 362 light treatment for, 578 Seasonal morphs, photoregulation of, 391–392 Seed germination See Germination Seedling establishment, photoreceptors in, 427–429 Seeds, light and habit of, 449–450 Self-association, 172–173 Sensory rhodopsins (SR), light energy use by, 204–205 Sesquiterpenes, as type I phototoxins, 482 Shade avoidance, by plants, 430–431, 448–449 Shift work, light significance in, 360–361 Short day plants (SDP) flowering in, 438–439 promoters and inhibitors, 446–447 genetic approaches to, 441–445 circadian rhythm mutants, 444–445 photoreceptors, 442–444 photoreceptors and circadian rhythm, 444 physiological approaches of, 439–441 Shot noise measurement, for very weak light measurement, 88 Index SID See Standard erythema dose Signaling, in photoperiodism, 445–447 genetic analysis of, 445–446 long-distance, 446–447 Signal transduction, of photoreceptors, 431–438 expression profiling, 436–437 mutants and interacting factors, 432–436 pharmacological approaches, 437–438 primary reactions, 431–432 Sinapates, in plants, 450 Single thin layer, reflection in, 178–183 Singlet oxygen configurations of, 37, 39 description of, 37 formation of, 37–38 Skeleton photoperiods, for photoperiodism and circadian rhythms, 397, 400–401 Sleep disorders, light treatment in, 361–362 Smog, UV-B creation of, 519 Smoking, photoageing and, 560 Snails, vitamin D in, 539 Snakes, vision problems and solutions of, 232 Snell’s law, 14 Snow cover reflection from, 127 UV-B and, 128 Solar energy collectors, nipple arrays and, 244 Sound waves, light waves v., Spectral absorption coefficient, of light underwater, 132 Spectral scattering coefficient, of light underwater, 132 Spectral tuning in biology, 155–189 anthocyanins, 171–177 chlorophyll a and b tuning, 161–162 chlorophyll absorption and molecular structure, 159–160 cyanobacterial phycobilisomes chromatic adaptation, 164–166 interplay of spectra in living world, 188–189 introduction to, 155–156 living mirrors and structural color tuning, 177–188 phycobiliproteins and phycobilisomes, 162–164 pigment spectra determination, 157–159 plant color, 156–157 visual tuning, 166–171 description of, 155 Spectrophotometry, light propagation in, 19 Spectroradiometry calibration of, 84–87 681 irradiance with improvised standard lamps, 85 irradiance with standard lamps, 84–85 irradiance with sunlight, 86–87 without standard lamp, 85 wavelength with daylight, 85–86 wavelength with lamps, 84 example of, 82–83 general, 80 input optics for, 80–82 for light measurement, 80–87 Specular reflection description of, 15 light propagation with, 19 Spherical aberration, 227–228 Spheroidene, absorption spectra of, 307–308 Sponges, bioluminescence of, 592 Spring parsley, phototoxicity of, 483 Squamous cell carcinoma (SCC) description of, 561 photoageing and risk of, 560 rates of, 561–562 risk of, 562–563 SR See Sensory rhodopsins Standard erythema dose (SID), MED v., 559 STED See Stimulated emission depletion microscopy Stefan-Boltzmann’s law, from Planck’s radiation law, 13 Stentorin, as light-sensing pigment, 197 Stepanov relationship, break down of, 31 Stimulated emission application of, 26 description of, 25–26 Jablonski diagram of, 26–27 Stimulated emission depletion microscopy (STED), 100–101 St John’s wort hypericin of, 482 phototoxicity of, 484 Stomata evolution of, 276–277 violaxanthin regulation of, 210–211 Stroma, carbon dioxide reduction in, 257–259 Structural colors living mirrors and tuning of, 177–188 introduction, 177–178 multilayer stack reflection, 183–188 single thin layer reflection, 178–183 pigments with, 187 in spectral tuning, 155 Sugar groups, in anthocyanins, 174–175 Sun, as light source, 51 Sunburn, of human skin, 558–559 682 Index Sunlight bright and dark sides of, 545–546, 577 modification of, 123–124 for psoriasis, 579–580 Suntan lotion, phototoxicity of, 485–486 Superposition eye, 236 Suprachiasmatic nucleus (SCN) in circadian rhythms, 352–353 inputs to, 354–355 intercellular communication in, 355 mammalian retina v., 358 overview of, 351 structure and function of, 351–353 Tapetum, mirror optics in, 241 Teaching experiments and demonstrations, 617–643 bioluminescence, 642 chloroplast pigment separation, 627–629 color and the benham disk, 622 complementary chromatic adaptation of cyanobacteria, 620–621 good start, 618–619 introduction, 617 light acclimation of leaves, 629–635 miscellaneous, 643 photomorphogenesis, 638–640 phytochrome spectrophotometric studies, 640–642 previtamin D photosynthesis, 624–625 protochlorophyllide photoconversion, 625–627 rhodopsin photoconversion, 623–624 singlet oxygen, 620 UVR damage and photoreactivation, 635–637 UVR damage to microorganisms, 637–638 wave nature of light, 619 Temperature absorption spectra differences with, 28–29 for Neurospora circadian system entrainment, 341–342 plant adaptation to, 450–451 Terrestrial daylight, 123–129 cloud effects on, 127 ground and vegetation effects on, 127 introduction to, 123 sunlight modification, 123–124 UV-A, visible, and infrared components of, 124–127 UV-B spectrum and biological action, 128–129 Tetracycline, phototoxicity of, 485–486 Tetrahydroporphin, molecular structure and absorption of, 159–160 Thermopile diagram of, 72 for light quantization, 42, 71–72 use of, 71 Thermopneumatic devices, for light measurement, 72–73 Thiophenes, as type II phototoxins, 482 Thylakoid membranes electron transporters in, 260 photosynthesis in, 257–258 Thymine, UVR transformation of, 511–512 Trans-cis isomerization, 198–211 archaean rhodopsin, 203–205 eukaryotic rhodopsin, 200–203 introduction to, 198–199 photosensor for chromatic adaptation of cyanobacteria, 209–210 phytochrome, 207–209 PYPs, 205–207 urocanic acid, 199–200 violaxanthin, 210–211 Transcription-translation oscillator (TTO), of Neurospora, 338–340 Transition moment, 31–33 Transmission gratings, in monochromators, 64 Transmission spectrum, for interference filter, 61–62 Transparency in biological world, 183, 187–188 with nipple arrays, 240–241 Trichromatic color vision, in humans, 168–169 Trilobites, compound eyes of, 235–236 Triplet state(s) description of, 35–36 of dioxygen molecule, 36–37 TTO See Transcription-translation oscillator Tumors, UVR immunosuppression and, 568 Two-photon excitation fluorescence microscopy, 99–100 UCA See Urocanic acid Ultradian rhythms, in organisms, 322 Ultraviolet A radiation (UV-A) apoptosis induction, 519 definition of, in photoallergic contact dermatitis, 572 photochemotherapy with, 581–582 phototherapy, for scleroderma, 580 in phototoxicity, 482 receptors for, 518 in terrestrial daylight, 124–127 UV-B v., 128 Index Ultraviolet B radiation (UV-B) apoptosis induction, 519 aquatic organisms influence of, 134 aquatic system influence of, 519–520 cellular redox equilibrium and, 567 in daylight, 504, 507 changes in, 507–510 definition of, in ecological context, 520–523 aquatic life, 520–522 terrestrial life, 522–523 filters for, 60 molecular effects of, 508–519 apoptosis induction, 519 on DNA, 508–512 on lipids, 516–517 photolyases and photoreactivation, 513–515 on proteins, 518 reactive oxygen species, 515–516 regulative processes, 518 ozone absorption of, 504 photoageing from, 560 phototherapy, for psoriasis, 578–580 in phototoxicity, 482 plant receptors for, 215–217 for previtamin D formation, 533 receptors for, 518 spectroradiometry for, 82 spectrum and biological action of, 128–129 sunburn from, 558 ultraweak light emission and, 609 underwater, 131 depth v., 135–137 vitamin D from, 532–533 Ultraviolet C radiation (UV-C) CFC decomposition by, 506–507 definition of, human eye effects of, 523–524 ozone creation by, 504 ultraweak light emission and, 609 Ultraviolet radiation (UVR) anthocyanin influence of, 175–176 aquatic organisms influence of, 134 bioluminescence protection from, 596 definition of, effects of demonstration for, 635–638 introduction to, 503–504 gas discharge lamp as source of, 54–55 HPV and, 569–570 HSV and, 569 683 human eye effects of, 523–525 human skin effect of, 557–558 immunosuppression, 565–568 immunosuppression and microbial infection, 568–570 immunosuppression and tumors, 568 on inanimate matter of biological relevance, 519–520 litter effect of, 520 microorganisms affect of, 144–146 for microscopy, 98 PAH absorption of, 490 plant avoidance of, 450 positive effects of, 545–546 receptors for, 518 ROS from, 515–516 skin cancer from, 553–554 vaccinations and, 570 vitamin D as defense for, 540 in water, 492 Ultraviolet vision chromatic aberration with, 231 evolutionary pressures for, 170 in insects, 176–177 in vertebrates, 168 Ultraweak light emission components of, 609 description of, 591 Underwater light, 131–137 apparent optical properties, 133–134 inherent optical properties, 132 introduction to, 131 in-water radiant energy estimation, 134–137 Urocanic acid (UCA) isomerization of, 199–200 mechanism of, 200 UVR absorption in skin, 565–566 Uropygial gland, vitamin D production in, 544 UV-A See Ultraviolet A radiation UV-B See Ultraviolet B radiation UV-C See Ultraviolet C radiation UVR See Ultraviolet radiation Vaccinations, UVR exposure and, 570 Velocity of light, 6–7 Vibrational energy electronic and rotational energy v., 23 in molecules, 23 Violaxanthin absorption spectra of, 307–308 as blue-light sensor in stomatal regulation, 210–211 description of, 211 684 Index Visible light phototherapy, for neonatal jaundice, 580–581 in terrestrial daylight, 124–127 UV-B v., 128 Vision spectral tuning in, 155–156 trichromatic color, 168–169 ultraviolet, 168 Visual tuning, 166–171 Vitamin A, in photoperiodism photoreceptors, 407 Vitamin D See also Previtamin D; Provitamin D biogeographical aspects of, 542–545 calcium regulation by, 538–540 cellular effects and receptor for, 537 chemistry and photochemistry of, 532–536 synthesis of, 532–533, 545, 554 evolutionary aspects of, 538–540 photobiological and ecological aspects of, 531–546 introduction, 531–532 plant kingdom distribution of, 540–541 population levels of, 545–546 regulation of, 542 transport and transformation of, in human body, 536 Vitamin D2 structure of, 532–533 vitamin D3 v., 538 Vitamin D3 structure of, 532–533 vitamin D2 v., 538 Water conservation of, 274–275 as light filter, 58–59 light under, 131–137 apparent optical properties, 133–134 inherent optical properties, 132 introduction to, 131 in-water radiant energy estimation, 134–137 UV-B, 129 oxidation of in first photosynthesis, 260 in photosynthesis, 257–258 plant adaptation to, 450–451 sunlight modification by, 123–124 UV-B effect on, 519–520 UVR exposure in, 492 Watt, 46 Wavelength absorption coefficient v., in water, 135 Balmer series for coordination of, 22 of daylight, 156 definition of, 6–7 in light-dependent magnetic compass, 467–469 for birds, 468–469 for newts, 467–468 light quantification problems with, 42–43 in light scattering, 18–19 refractive index v., 15, 230 scalar irradiance v., in water, 135–136 spectroradiometer calibration for with daylight, 85–86 with lamps, 84 Wave properties, of light, 1–6 definitions, 6–7 Weighting functions, for "sunburn" meters, 48–49 Whales, vision of, 171 Wien’s law, from Planck’s radiation law, 13 Xanthophyll cycle evolution of, 269 introduction to, 630–633 spectral tuning and, 158–159 Xenon lamps, 54–55 Xeroderma pigmentosum (XP) as genodermatoses, 570 NMSC and, 562 XP See Xeroderma pigmentosum YAG laser, 57 Young’s double slit experiment description of, 1–3 diffraction in, 7–8 with photocell, 2–6 Zeaxanthin chlorophyll a quenching by, 313–314 in stomatal regulation, 211 ... part of the drawing the same is shown for a circularly polarized beam the light becomes partially plane-polarized Skylight is a mixture of circularly and plane-polarized light, which we call elliptically... sum of equally strong left- and right-handed components of circularly polarized light Natural light, such as direct sunlight, is often almost unpolarized, i.e., a random mixture of all possible...Photobiology Second Edition Photobiology The Science of Life and Light Second Edition Edited by Lars Olof Björn Lund University Lund, Sweden Lars Olof Björn Department of Plant Physiology Lund University