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Index Absorbance, 34 Acridine orange, 87 Action spectra bacteria killing, 43 carotenogenesis, 294 circadian rhythm, 201 conidiation, 297 defined, 42 erythema (skin), 159 extraretinal photoreception, 220 germination, 278 inhibition of hypocotyl elongation, 282 mutagenesis, 126 photoaccumulation, 326 photokinesis, 319 photophobic response, 319, 329 phototaxis, 319, 326 positive phototropic curvature (oat), 294 prevention of flowering, 278 retardation of cleavage, 44 reversing germination, 278 vision, 3, 258 (6-4)-Adducts, 71, 128 Aequorin, 401, 416 Anaerobic photosensitization, 84 AP endonuclease, 123, 124 Aquatic ecosystems, 140 Aschoff's rule, 223 Auxotroph, 128 Beer's law, 34, 55, 114, 142 Bilirubin, 183 Bimolecular reaction, 52 Biological chemiluminescence, 413 Biological clock, 208 Biological consequences of stratospheric ozone depletion animals, 151 plants, 148 Bioluminescence applications of, 414 bacteria, 409 Bioluminescence (cont' d) biological chemiluminescence, 413 bioluminescent organisms, 392 systems, 397 chemiluminescence, 397 CIEEL mechanism, 399,402,411, 412 coelenterate, 404 Cypridina, 404 firefly, 401 gene transfer, 416 generalizations about, 400 light organs, 393 mechanism, 396 occurrence, 392 origin and function, 394 Blackbody radiation, 6 Blue-light mediated morphogenesis events under blue-light control, 292 photoreceptor(s), 294 Carbonyl compounds, 54 Carcinogenesis (skin), 166 [3-Carotene, 186, 338, 352 Carotinoids, 353 Chemical restoration, 119 Chemiluminescence, 397,413 Chlorophyll, 87, 351 Chloroplast, 333, 375 Chromophore (defined), 5 CIEEL mechanism, 399,402,411,412 Circadian rhythms action spectra, 201 Aplysia,218 Aschoff's rule, 223 biochemical rhythms, 202 cellular mechanisms, 201 clock mutants, 206 defmed,196 drug sensitivity, 210 entrainment to artificial light, 194 419

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Page 1: The Science of Photobiology ||

Index

Absorbance, 34 Acridine orange, 87 Action spectra

bacteria killing, 43 carotenogenesis, 294 circadian rhythm, 201 conidiation, 297 defined, 42 erythema (skin), 159 extraretinal photoreception, 220 germination, 278 inhibition of hypocotyl elongation, 282 mutagenesis, 126 photoaccumulation, 326 photokinesis, 319 photophobic response, 319, 329 phototaxis, 319, 326 positive phototropic curvature (oat), 294 prevention of flowering, 278 retardation of cleavage, 44 reversing germination, 278 vision, 3, 258

(6-4)-Adducts, 71, 128 Aequorin, 401, 416 Anaerobic photosensitization, 84 AP endonuclease, 123, 124 Aquatic ecosystems, 140 Aschoff's rule, 223 Auxotroph, 128

Beer's law, 34, 55, 114, 142 Bilirubin, 183 Bimolecular reaction, 52 Biological chemiluminescence, 413 Biological clock, 208 Biological consequences of stratospheric ozone

depletion animals, 151 plants, 148

Bioluminescence applications of, 414 bacteria, 409

Bioluminescence (cont' d)

biological chemiluminescence, 413 bioluminescent

organisms, 392 systems, 397

chemiluminescence, 397 CIEEL mechanism, 399,402,411, 412 coelenterate, 404 Cypridina, 404 firefly, 401 gene transfer, 416 generalizations about, 400 light organs, 393 mechanism, 396 occurrence, 392 origin and function, 394

Blackbody radiation, 6 Blue-light mediated morphogenesis

events under blue-light control, 292 photoreceptor( s), 294

Carbonyl compounds, 54 Carcinogenesis (skin), 166 [3-Carotene, 186, 338, 352 Carotinoids, 353 Chemical restoration, 119 Chemiluminescence, 397,413 Chlorophyll, 87, 351 Chloroplast, 333, 375 Chromophore (defined), 5 CIEEL mechanism, 399,402,411,412 Circadian rhythms

action spectra, 201 Aplysia,218 Aschoff's rule, 223 biochemical rhythms, 202 cellular mechanisms, 201 clock mutants, 206 defmed,196 drug sensitivity, 210 entrainment to

artificial light, 194

419

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420

Circadian rhythms (cont'd) entrainment to (cont' d)

environmental cycles, 197 free-running rhythms, 195 general features, 194 genetic control, 204 house sparrow, 223 humans, 208 inhibitors of, 203 jet lag, 208 location of circadian clocks, 206 locomotry activity, 195, 221 melatonin production, 218 organization in multicellular organisms, 206 photoreceptor for, 200 pineal gland, 218 sensitivity of mice to endotoxin, 210 stopping the clock with bright light, 224 temperature compensation, 196

Circular dichroism (CD), 40 Clock (circadian), 196, 206, 221 Crassulacean acid metabolism, 379 Cyclobutane-type dimers

absorption spectrum, 69 defmition, 68 monomerization, 68 structure, 49, 69

Cysteine, 49 5-S-Cysteinyl-6-hydrothymine, 49 Cytochrome, 367, 371

Do, 114, 115 D37 , 114, 115 Dq ,115 DNA-DNA cross-links, 72 DNA photochemistry

DNA-DNA cross-links, 72 DNA-protein cross-links, 49, 68, 70, 72 strand breaks, 72

DNA-protein cross-links, 49, 68, 70, 72 DNA repair

daughter-strand gaps, 124 double-strand breaks, 124 excision repair, 120 genes of E. coli, 117 mismatch repair, 130 photoreactivation, 119 postreplication repair, 123 skin (human), 162 spontaneous mutagenesis, 130 UV radiation mutagenesis, 126

DNA replication mismatch repair, 130 spontaneous mutagenesis, 130 UV radiation mutagenesis, 127

DNA strand breaks, 72 7-Dehydrocholesterol, 51 Dihydrocytosine, 70 Dihydrothymine, 68 Dipole-dipole interaction, 32 Dipole moment, 29 Dose modification factor, 116

Early receptor potential, 259 Electromagnetic

spectrum, 3 theory of Maxwell, 4

Electron spin, 25 Electroretinogram, 258 Energy transfer

dipole-dipole, 32 Forster-type, 32, 60 resonance excitation, 32 triplet-triplet, 33

Entrainment (circadian rhythms), 194, 197 Environmental photobiology

aquatic ecosystems, 140

Index

biological consequences of stratospheric ozone depletion, 147

biologically effective UV-B, 139 evolution of terrestrial organisms, 138 protection from solar UV radiation

animals, 145 aquatic organisms, 145 terrestrial plants, 143

solar radiation, 136 stratospheric ozone, 138 vegetation canopies, 141

Enzyme inactivation kinetics, 64 theories, 62

Eosin Y, 87 Error-prone DNA repair, 127, 128 Evolution

stratospheric ozone (effects on), 138 terrestrial organisms, 138

Extrapolation number, 115 Excision repair

base excision, 123 nucleotide excision, 120

Excited states atomic orbitals, 27 energy levels, 27 molecular orbitals, 26 pi orbital, 27, 54 production, 25 sigma orbital, 27

Extraretinal photoreception Aschoff's rule, 223 circadian rhythms, 218, 221, 223

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Index

Extraretinal photoreception (cont' d) house sparrow, 221 invertebrates, 217 mammals, 227 photoperiodism, 224 reproductive system, 224 sites for, 227 vertebrates, 218

Eye light damage, 266 responses to light, 255 role in circadian rhythms, 222 structure and function, 231 UV radiation effects on, 189 vision, 231ff.

Ferredoxin, 367 Finsen (Neils), 179 Firefly, 401 First law of photochemistry, 5, 47, 80 First-order reaction, 31, 54 Flash photolysis, 35, 52, 58, 59, 61, 62 F1avinoid, 137, 140, 144, 296 Fluence, 54 Fluorescence

defined, 31 depolarization, 40 lifetime, 38 polarization, 39 quenching, 32 spectra, 36

F1uorophore (defined), 32 5-Formyluracil, 68 Forster mechanism, 32, 52, 60, 356 Fovea, 234, 238 Frequency doubling, 11 Furanocoumarin, 84, 104

Genetic control circadian rhythms, 204 DNA repair, 117

Glycosylases, 118, 123, 124 Goeckerman regimen, 181 Grana, 375

Hematoporphyrin, 87, 182 Herpes simplex infection, 182 Heteroaddition reactions, 66 Humans

circadian rhythms, 208 DNA repair, 162 jet lag, 208 photomedicine, 155ff. phototherapy, 179

Humans (cont'd) skin [see Skin (human)] sunburn, 158

Hydrated electron, 58, 60, 61 6-Hydroxy-5-hydrouracil, 50, 67 5-Hydroxymethyl uracil, 68 8-Hydroxypsoralen, 88 Hypericin, 87 Hypersensitivity, 187

Inducible responses adaptive response, 118 heat shock system, 119 oxidation system, 119 SOS system, 117

Internal conversion, 30 Intersystem crossing, 30 Isopsoralen, 88

Jablonski diagram, 30, 51 Jaundice (neonatal), 183 Jet lag, 208

Lambert-Bouguer-Beer law, 34, 55, 114, 142 Lamella, 375 Lasers

average power, 13 defined, 10 frequency doubling, 11 medical uses, 24 mode-locking, 13 peak power, 13 properties of, 11 Q switching, 13 schematic diagram, 11

Laws of photochemistry first, 5, 47, 80 second,399

LIAC, 294 Lifetimes

singlet state, 52 triplet state, 52

Light-induced absorbance change (LIAC), 294 Light scattering, 22 Light sources

arc lamps, 6, 7 fluorescent lamps, 6, 8 incandescent lamps, 6 lasers, 6, 10 photochemical lamps, 6

Light-tissue interactions, 22 Liquid-holding recovery, 75 Luciferase, 396 Luciferin, 396 Lumazine protein, 411

421

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422

Lumiflavin, 50 Lupus erythematosus, 185

Melatonin production, 218 Melanogenesis, 163 Methionine, 84 8-Methylpsoralen, 88 Methyltransferase, 1I8 5-Methyoxypsoralen, 186 8-Methoxypsoralen, 88, 181, 186 Methylene blue, 87 Mismatch DNA repair, 130 Molar extinction coefficient

amino acids, 63 defined, 34, 56, 63 peptide bonds, 63

Molecular orbitals defined, 26 ethylene, 28, 29 formaldehyde, 28, 29

Mutagenesis assays, 127 errors in

DNA repair, 127 DNA replication, 127

genetic control, 129 kinetics, 128 mismatch repair, 30 postreplication repair, 124 spontaneous, 130 types of mutations, 127 UV radiation, I 26ff.

Mutations (types of) back, 128 base substitution, 127 frameshift, 127 missense, 127 nonsense, 127 transition, 127 transversion, 127 suppressor, 128

n (extrapolation number), 115 Near-UV radiation, 75, 1I9

Optical activity, 40 Optical components

filters, 15 lenses, 13 mirrors, 13 monochromators, 15 optical fibers, 15 polarizers, 18

Optical density, 34 Optical rotatory dispersion (ORO), 40 Ozone (see Stratospheric ozone)

Pauli exclusion principle, 25 Phosphorescence

defmed,31 spectra, 36

Photoaccumulation, 310 Photoacoustic spectroscopy, 41 Photochemistry

amino acids, 58, 63 bimolecular reaction, 52 carbonyl compounds, 54 chain reactions, 54 cholesterol, 50 cycloaddition, 49 7-dehydrocholesterol, 51 deoxyribose, 66 DNA,72 fundamentals, 47 isomerization, 51, 183, 185 kinetics, 54 linear addition, 49 nucleic acids, 65 overall reactions, 49 photofragmentation, 50 photohydration, 50, 66 photooxidation, 50 photorearrangement, 51 polymerization, 54 primary reactions, 51 proteins, 57, 60 purines, 66 pyrimidines, 66 quantitation, 75 retinal, 51 riboflavin, 50 RNA,74 unimolecular reaction, 52 uracil, 50

Photochromic adaptation, 298 Photodetectors

properties of, 21 quantum, 20 thermal, 20

Photodinesis, 310, 322 Photodispersal, 310 Photodynamic action, 80 Photohydration, 50, 66 Photoimmunology

delayed hypersensitivity, 187 immediate hypersensitivity, 187

Photokinesis, 308 Photon (defined), 4 Photomedicine

effects of sunlight on the eye, 189 effects of sunlight on skin

acute, 158

Index

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Index

Photomedicine (cont' d) effects of sunlight on skin (cont' d)

adverse, 171 beneficial, 178 chronic, 164

optical properties of skin, 156 photoimmunology, 187 photoprotection, 184 phototherapy, 179

Photometric units, 12 Photomorphogenesis

blue-light mediated, 291 diversity of, 274 light as a source of information, 273 photochromic adaptation, 298 phytochrome mediated, (see Phytochrome) UV-radiation mediated, 296

Photomovement cells, 315, 318 Chlamydomonas, 325 chloroplasts, 333 cyanobacteria, 319 ecological significance, 341 Halobacterium halobium, 323 methods, 311 organelles, 332 photoaccumulation, 310 photodinesis, 310, 322 photodispersal, 310 photokinesis, 308 photophobic response

defined, 309 enhancers, 331 inhibitors, 331

photosensory transduction mechanisms, 317 phototaxis, 309 phototropism, 310, 316, 337 polarotropism, 310, 333 primary photoreactions, 320, 324, 328, 330 terminology, 307 Stentor coeruleus, 329

Photoperiodism, 224 Photophobic response, 309 Photoprotection, 75 Photoreception

extraretinal, 215 phototropism, 337 vision, 231 ff.

Photoresponse (defined), 308 Photosensitization

amino acids, 90 anaerobic reactions, 84 animals, 98 biological applications of, 101 cells, 93

Photosensitization (cont'd) evolution and, 104 humans, 100 lipids, 88 mechanisms (diagnostic techniques), 85 natural photosensitivity, 10 I nucleic acids and derivatives, 89 photocarcinogenesis, 100 photodynamic action, 80 plants, 97 proteins, 90 radicals, 82 singlet oxygen, 83, 85 type I (radical) mechanism, 82 type II (singlet oxygen) mechanism, 83 types of photosensitizers, 86-88 viruses, 92

Photosynthesis accessory pigments, 354 anoxygenic, 359 antenna molecules, 356 as an energy source, 380 brown algae, 354 C3 carbon cycle, 377 C4 carbon cycle, 377 carbon dioxide fixation, 377 cyanobacteria, 354 cytochrome, 371 delayed fluorescence, 357 diatoms, 354 dinoflagellates, 354 electron transport pathways, 361, 365 green algae, 355 green bacteria, 353, 361, 362 higher plants, 355 light harvesting, 350, 355 overview, 349 ' oxygen evolution, 371 oxygenic, 365 P680, 369 pheophytin, 369 phosphorylation, 373 photochemical conversion, 355 photorespiration, 379 photosynthetic unit, 358 photosystem I, 365 photosystem II, 368 pigments, 351 plastoquinone, 370 purple bacteria, 353, 359 QA and QB, 370 reaction center, 357 red algae, 354 structural organization, 375 Z-scheme, 365

423

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424

Photosynthetically active radiation (PAR), 138 Phototaxis, 309 Phototechnology

actinometry, 21 light sources, 5 light-tissue interactions, 22 optical components, 13 photodetectors, 20 radiometry, 21

Phototherapy, 179 Phototropism, 310, 316, 337 Phycoerythrobilin, 352 Phycomyces, 316, 337 Phytochrome

differences between P, and Prp 288 discovery, 278 events under phytochrome control, 280 mode of action, 290 photochemical reactions, 286 photoequilibria, 283 properties of, 285

Pineal gland, 188, 218 Polarotropism, 310, 333 Polarization, 18, 39 Polymorphous light eruptions, 185 Primary photochemical reactions

excited state electron transfer, 53 hydrogen atom abstraction, 54 photoionization, 53 production of free radicals, 53 singlet-singlet energy transfer, 52 triplet-triplet energy transfer, 52

Prototroph, 128 Psoralen, 84, 88, 181, 186 Psoriasis

description of, 180 Goeckerman regimen, 181 PUVA therapy, 181, 186

Pulse radiolysis, 58 PUVA, 181, 186 Pyronine Y, 87

Quanta (defined), 4 Quantum mechanics, 27 Quantum yield

amino acids, 63 bioluminescence, 399 defmed, 31, 48, 63 inactivation, 114 proteins, 64 thymine, 68 uracil, 68

Quenching fluorescence, 32 singlet oxygen, 85

Quick-tarming preparations, 186

Race tube culture, 200 Radiative transitions, 25 Radicals, 59, 82 Radiometric units, 12 Recovery phenomena

liqnid holding recovery, 115 medium-dependent resistance, 116 minimal medium recovery, 116 plateau phase recovery, 116

Resonance excitation transfer, 32 Retina

light damage to, 266 structure, 231, 236

Retinal, 51 Rhodopsin, 244, 245, 249, 252, 254, 255 Riboflavin, 50, 87 Rose bengal, 87

Second law of photochemistry, 399 Second order reaction, 54 Shuttle vectors, 128 Singlet oxygen

defined, 83 lifetime, 85 light emission, 85 quenching, 85 trapping, 85 type II mechanism, 83

SI units, 2 Skin (human)

carcinogenesis, 166 diseases, 179 DNA repair, 162 effects of sunlight

acute, 158 adverse, 171 beneficial, 178 chronic, 164

melanogenesis, 163 microscopic anatomy, 160 optical properties, 156 photosensitizing agents, 185 quick-tanning preparations, 186 structure, 157 sunburn, 158

Solar urticaria, 185 Solar radiation

biological effectiveness, 138 effects on humans, 155ff. environmental photobiology, 135ff. filtration by vegetation canopies, 141 how organisms protect themselves

animals, 143 aquatic organisms, 143 plants, 143

penetration into water, 140

Indel

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Index

Solar radiation (cont'd) photomedicine, 155ff. spectrum, 3, 136 transmission through leaf epidennis, 149

Spectrophotometers absorption, 35 luminescence, 37

Spectroscopy absorption, 34 action, 42 Lambert-Bouguer-Beer law, 34 luminescence, 35 optical activity, 40 photoacoustic, 41

Spectrum action, 42 atomic, 27 benzyl radical, 59 bioluminescence, 395, 411, 412 black body radiation, 6 carbazole, 36 a-chymotrypsin, 61 deuterium lamp, 7 electromagnetic, 3 epidennal transmittance, 137 filter transmission, 16 flavin (nuclear magnetic resonance), 410 flavinoid, 137 fluorescent lamps, 9 gennicidal lamp, 8 green fluorescent protein from Renilla, 408 halogen lamp, 7 hydrated electron, 59 mercury lamp, 7 nucleic acid, 137 ozone, 137 phenylalanine, 57 photoacoustic, 41 photoflood lamp, 3 photopic vision, 3 photosynthetic organisms, 353 protein, 137 retinal (II-cis), 249 rhodopsin, 249 Schiff bases, 248 shadelight (under a broadleaf canopy), 283 sunlight, 3, 137, 283 thymine, 69 thymine dimer, 69 tryptophan, 57 tryptophan radical, 59 tyrosine, 57 tyrosine radical, 59 xenon lamp, 7

Speed of light, 4 Spontaneous mutagenesis, 130

Spore photoproduct, 70 Stokes shift, 36 Stratospheric ozone

depletion (biological consequences), 147 fonnation, 138 seasonal variation, 138 spectrum, 173

Stroma, 375 Suicide enzyme, 118 Sunburn, 158 Sunscreens, 184 Superoxide radical anion, 58 Survival curves

alpha-beta model, 114 dose modification factor, 116 E. coli B, 112 E. coli K-12, 113 parameters, 115 target theory, 114

Target theory, 114 Tautomeric shifts (DNA bases), 130 Thiopyronin, 87 4-Thiouridine, 75 Thymine, 49 Thymine dimer, 49 Thymine glycol, 67 5-Thyminyl-5,6-dihydrothymine, 70

425

Time scales in photophysics, photochemistry, and photobiology, 48

Transition dipole, 30 moment, 29

Translesion DNA synthesis, 129 Transmittance, 34 Triplet-triplet

annihilation, 53 energy transfer, 70

Tumor therapy, 182 Type r (radical) reaction, 82 Type II (singlet oxygen) reaction, 83

Ubiquinone, 361 Unimolecular reaction, 52 Units

photometric, 12 radiometric, 12 Sr,2

Uracil, 50 UV-A, 5, 59, 80, 84, 149, 156,296 UV-B, 5, 80, 137, 138, 156,296 UV-C, 5, 51, 80 UV radiation effects

animals, 145 beneficial effects, 51, 178 destruction of oxygen evolution by plants, 372

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426

UV radiation effects (cont'd) DNA repair, 119 environmental photobiology, I 35ff. eyes, 189 humans, I 55ff. inducible responses, 117 mutagenesis, 126 photomorphogenesis, 296 plants, 143 recovery phenomena, 115 skin, 155 solar spectrum, 3, 136 survival curves, 112

UV -radition-mediated morphogenesis events under UV radiation control, 296 photoreceptor(s), 297

Vibrational relaxation, 30 Vision

bleaching intermediates of rhodopsin, 255 calcium model, 263 cGMP model, 264 cones, 237, 253 dichroism of outer segments, 250 disk membrane, 240, 243, 250 early receptor potential, 259 electroretinogram, 258 eye

light damage, 266 responses to light, 255 structure and function, 223

eye (cont'd) fovea, 234, 238 internal transmitter hypothesis, 262 mechanism of visual excitation, 262 overview, 231 patch clamp, 262 recording of photoreceptor responses, 260 retina, 231, 236, 266 rhodopsin, 244, 245, 249, 252, 254, 255 rods, 237, 253, 262 rod outer segments (ROS), 241 visual cycle, 257 visual pigments, 246, 248, 251 vitamin A, 243, 246

Vitamin A, 243, 246 Vitamin 0 3 ,51, 178 Vitiligo, 179

Xanthophylls, 354

Index

Xeroderma pigmentosum, 122, 163, 166-168, 170 X radiation

chemical restoration, 119 diffraction (rod outer segment), 243 medium-dependent resistance, 116 mutagenesis, 128, 129 plateau phase recovery, 116 repair of

DNA double-strand breaks,. 126 potentially lethal damage, 116

Z-scheme, 365