Ultraviolet radiation screening compounds

Biological Reviews

Volumn 74, Issue 3, 1999, Pages 311-345

  Cockell, Charles S ^a   Knowland, John ^b  

^a University of Oxford ^*

^b UNIVERSITY OF OXFORD   (United Kingdom)

Author keywords

Compounds; Evolution; Screening; UV radiation

Indexed keywords

CHEMICAL COMPOUND; OZONE;

ANIMAL; BIOSPHERE; ECOLOGY; EVOLUTION; MICROORGANISM; NONHUMAN; PHYSIOLOGY; PLANT; RADIATION INJURY; RADIATION PROTECTION; REVIEW; ULTRAVIOLET RADIATION;

AMINO ACIDS; ANIMALS; EVOLUTION; HUMANS; INDOLES; MELANINS; PHENOLS; PIGMENTS, BIOLOGICAL; PLANT PHYSIOLOGY; RADIATION INJURIES; RADIATION-PROTECTIVE AGENTS; ULTRAVIOLET RAYS;

ANIMALIA;

EID: 0032800184     PISSN: 14647931     EISSN: 1469185X     Source Type: Journal    
DOI: 10.1111/j.1469-185X.1999.tb00189.x     Document Type: Article

References (242)

1. Adams, N. L., Carroll, A. K. & Shick, J. M. (1996). Mycosporine-like amino acids provide protection against ultraviolet radiation in eggs of the green sea urchin Strongylocentrus droebachiensis. Photochemistry and Photobiology 64, 149-158.
2. Adams, W. W., Demmig-Adams, B. & Lange, O. L. (1993). Carotenoid composition and metabolism in green and blue-green algal lichens in the field. Oecologia 94, 576-584.
3. Aikin, A. C., Chandra, S. & Stecher, T. P. (1980). Supernovae effects on the terrestrial atmosphere. Planetary and Space Science 28, 639-644.
4. Algra, R. J. & Knox, J. M. (1978). Topical photoprotective agents. International Journal of Dermatology 17, 628-634.
5. Allen, J. M., Gossett, C. J. & Allen, S. K. (1996). Photochemical formation of singlet molecular oxygen in illuminated aqueous solutions of p-aminobenzoic acid (PABA). Journal of Photochemistry and Photobiology B: Biology 32, 33-37.
6. Ananthaswamy, H., Loughlin, M., Cox, P., Evans, R. L., Ullrich, S. E. & Kripke, M. L. (1997). Sunlight and skin cancer: Inhibition of p53 mutations in UV-irradiated mouse skin by sunscreens. Nature Medicine 3, 510-514.
7. Arancibia, A., Borie, G., Cornwell, E. & Medrano, C. (1981). Pharmacokinetic study on the percutaneous absorption of p-aminobenzoic acid from three sunscreen preparations. Il Farmaco Ed. Prat. 36, 357-365.
8. 14C-acetate, studied in synchronous cultures. Planta 107, 1-32.
9. Atsatt, P. R. (1988). Are vascular plants ‘inside-out’ lichens Ecology 69, 17-23.
10. Avery, J. A., Bowmaker, J. K., Djamgoz, M. B. A. & Doning, J. E. G. (1983). Ultraviolet sensitive receptor in a freshwater fish. Journal of Physiology 334, 23-24.
11. Bachereau, F. & Asta, J. (1997). Effects of solar ultraviolet radiation at high altitude on the physiology and the biochemistry of a terricolous lichen (Cetraria islandica L.). Symbiosis 23, 197-217.
12. Bahnemann, D. F. (1994). Photocatalytic treatment of waters. In Aquatic and Surface Photochemistry (eds D. Crosby, G. Helz and R. Zepp), pp. 261-316. Lewis Publishers, Boca Raton, FL.
13. Balch, W. M. & Haxo, F. T. (1984). Spectral properties of Noctiluca militaris Suriray, a heterotrophic dinoflagellate. Journal of Plankton Research 6, 515-525.
14. Barnes, P. W., Jordan, P. W., Gold, W. G., Flint, S. D. & Caldwell, M. M. (1988). Competition, morphology and canopy structure in wheat (Triticum aestivum L.) and wild oat (Avena fatua L.) exposed to enhanced ultraviolet-B radiation. Functional Ecology 2, 319-330.
15. Bebout, B. M. & Garcia-Pichel, F. (1995). UV-B induced vertical migrations of cyanobacteria in a microbial mat. Applied and Environmental Microbiology 61, 4215-4222.
16. Beggs, C. J., Stolzer-Jehle, A. & Wellmann, E. (1985). Isoflavanoid formation as an indicator of UV stress in bean (Phaseolus vulgaris L.) leaves. The significance of photorepair in assessing potential damage by increased solar UV-B radiation. Plant Physiology 79, 630-634.
17. Beggs, C. J. & Wellmann, E. (1985). Analysis of light-controlled anthocyanin formation in coleoptiles of Zea mays L.: The role of UV-B, blue, red and far-red light. Photochemistry and Photobiology 41, 481-485.
18. Berkner, L. V. & Marshall, L. C. (1965). History of major atmospheric components. Proceedings of the National Academy of Sciences 53, 1215-1225.
19. Blanc, P. L., Tuveson, R. W. & Sargent, M. L. (1976). Inactivation of carotenoid-producing and albino strains of Neurospora crassa by visible light, blacklight, and ultraviolet radiation. Journal of Bacteriology 125, 616-625.
20. Blank, I. H., Cohen, III J. H., Anderson, R. R., Jaenicke, K. F. & Parrish, J. A. (1982). Mechanisms of the protective action of sunscreens. Journal of Investigation Dermatology 78, 381-385.
21. Blum, H. (1961). Does the melanin pigment of human skin have adaptive role. Quarterly Reviews of Biology 36, 50-63.
22. Bohm, G. A., Pfeiderer, W., Boger, P. & Scherer, S. (1995). Structure of a novel oligosaccharide-mycosporine-amino acid ultraviolet A/B sunscreen pigment from the terrestrial cyanobacterium Nostoc commune. Journal of Biological Chemistry 270, 8536-8539.
23. Bonin, A. M., Arlauskas, A. P., Angus, D. S. W., Baker, R. S. U., Gallagher, C. H., Greenoak, G., Lane Brown, M. M., Meher-Homji, K. M. & Reeve, V. (1982). UV-absorbing and other sun-protecting substances: Genotoxicity of 2-ethylhexyl P-methoxycinnamate. Mutation Research 105, 303-308.
24. Booth, C. R. & Morrow, J. R. (1997). The penetration of UV into natural waters. Photochemistry and Photobiology 65, 254-257.
25. Bowmaker, J. K. & Kunz, Y. W. (1987). Ultraviolet receptors, tetrachromatic color vision and retinal mosaics in the brown trout (Salmo trutta): Age-dependent changes. Vision Research 27, 2101-2108.
26. Braun, J. (1991). The protective function of phenolic compounds of rye and oat seedlings against UV-B radiation and their biosynthetic regulation. Karls. Beitr. Entw. Okophysiol (M. Tevini, Ed.) 9, 1-237.
27. Braun, J. & Tevini, M. (1993). Regulation of UV-protective pigment synthesis in the epidermal layer of rye seedlings. Photochemistry and Photobiology 57, 318-323.
28. Britton, G. & Goodwin, T. W. (1981). Carotenoid Chemistry and Biochemistry. Pergamon Press, Oxford.
29. Brook, P. J. (1981). Protective function of an ultraviolet absorbing compound associated with conidia of Glomerella cingulata. New Zealand Journal of Botany 19, 299-304.
30. Budel, B., Karsten, U. & Garcia-Pichel, F. (1997). Ultraviolet-absorbing scytonemin and mycosporine-like amino acid derivatives in exposed, rock-inhabiting cyanobacterial lichens. Oecologia 112, 165-172.
31. Caldwell, M. M., Robberecht, R. & Flint, S. D. (1983). Internal filters: Prospects for UV-acclimation in higher plants. Physiologia Plantarum 58, 445-450.
32. Canuto, V. M., Levine, J. S., Augustsson, T. R. & Imhoff, C. L. (1982). UV radiation from the young Sun and oxygen and ozone levels in the prebiological paleoatmosphere. Nature 296, 816-820.
33. Carreto, J. I., Carignan, M. O., Daleo, G. & De Marco, S. G. (1990). Occurrence of mycosporine-like amino acids in the red-tide dinoflagellate Alexandrium excavatum-UV-photo-protective compounds Journal of Plankton Research 12, 909-921.
34. Carreto, J. I., De Marco, S. G. & Lutz, V. A. (1989). UV-absorbing pigments in the dinoflagellates Alexandrium excavatum and Procentrum micans. Effects of light intensity. In Red Tides Biology, Environmental Science and Toxicology (eds T. Okaichi, D. M. Anderson and T. Nemoto), pp. 333-336. Elsevier, New York.
35. Carroll, A. K. & Shick, J. M. (1996). Dietary accumulation of UV-absorbing mycosporine-like amino acids (MAAs) by the green sea urchin (Strongylocentrus droebachiensis). Marine Biology 124, 561-569.
36. Cerda-Olmedo, E., Martin-Rojas, V. & Cubero, B. (1996). Causes of cell death following ultraviolet B and C exposures and the role of carotenes. Photochemistry and Photobiology 64, 547-551.
37. Chioccara, F., Della Gala, A., De Ross, M., Novellino, E. & Prota, G. (1980). Mycosporine aminoacids and related compounds from the eggs of fishes. Bulletin Socíeté Chimique Belgique 89, 1101-1106.
38. Chioccara, F., Miscuraca, G., Novellino, E. & Prota, G. (1979). Occurrence of two new mycosporine-like amino acids, mytilins a and b in the edible mussel, Mytilus galloprovincialis. Tetrahedron Letters 34, 3181-3182.
39. Chioccara, F., Zeuli, L. & Novellino, L. E. (1986). Occurrence of mycosporine related compounds in sea urchin eggs. Comparative Biochemistry and Physiology 85 B, 459-461.
40. Chyba, C. & Sagan, C. (1992). Endogenous production, exogenous delivery and impact-shock synthesis of organic molecules: An inventory for the origins of life. Nature 355, 125-132.
41. Cleaves, H. J. & Miller, S. L. (1998). Oceanic protection of prebiotic organic compounds from UV radiation. Proceedings of the National Academy of Sciences, USA 95, 7260-7263.
42. Cockell, C. S. (1998a). The biological effects of UV radiation on early earth - a theoretical evaluation. Journal of Theoretical Biology, 193, 719-731.
43. Cockell, C. S. (1998b). UV radiation, evolution and the π-electron system. Biological Journal of the Linnean Society, 63, 449-457.
44. Cockell, C. S. (1999). Crises and extinction in the fossil record - a role for ultraviolet radiation Paleobiology 25, 212-225.
45. Craig, C. L. & Bernard, G. D. (1990). Insect attraction to ultraviolet-reflecting spider webs and web decorations. Ecology 71, 616-623.
46. Crippa, P. R., Cristofoletti, V. & Romeo, N. (1978). A band model for melanin deduced from optical absorption and photoconductivity experiments. Biochimica et Biophysica Acta 538, 164-170.
47. Crutzen, P. J. & Bruhl, C. (1996). Mass extinctions and supernova explosions. Proceedings of the National Academy of Sciences, USA 93, 1582-1584.
48. Cullen, J. C. & Neale, P. J. (1994). Ultraviolet radiation, ozone depletion, and marine photosynthesis. Photosynthesis Research 39, 303-320.
49. Davidson, A. T., Bramich, D., Marchant, H. J. & McMinn, A. (1994). Effects of UV-B irradiation on growth and survival of antarctic marine diatoms. Marine Biology 119, 507-515.
50. Davidson, A. T. & Marchant, H. J. (1994). The impact of ultraviolet radiation on Phaeocystis and selected species of antarctic marine diatoms. In Antarctic Radiation in Antarctica: Measurements and Biological Effects (eds C. S. Weiler and P. A. Penhale). Antarctic Research Series 62, 187-205.
51. Day, T. E. (1993). Relating UV-B radiation screening effectiveness of foliage to absorbing-compound concentration and anatomical characteristics in a diverse group of plants. Oecologia 95, 542-550.
52. Deamer, D. W., Mahon, E. H. & Bosco, G. (1994). Self-assembly and function of primitive membrane structures. In Early Life on Earth (ed. S. Bengtson), pp. 107-123. Columbia University Press, New York.
53. Delwiche, C. F., Graham, L. E. & Thomson, N. (1989). Lignin-like compounds and sporopollenin in Coleochaete, an algal model for land plant ancestry. Science 245, 399-401.
54. Demmig-Adams, B. & Adams, W. W. (1992). Photoprotection and other responses of plants to high light stress. Annual Review of Plant Physiology and Plant Molecular Biology 43, 599-626.
55. Donkor, V. A., Amewowor, D. H. A. K. & Häder, D.-P. (1993a). Effects of tropical solar radiation on the motility of filamentous cyanobacteria. FEMS Microbiology and Ecology 12, 143-148.
56. Donkor, V. A., Amewowor, D. H. A. K. & Häder, D.-P. (1993b). Effects of tropical solar radiation on the velocity and photophobic behavior of filamentous gliding cyanobacteria. Acta Protozoologica 32, 67-72.
57. Donkor, V. A. & Häder, D.-P. (1995). Protective strategies of several cyanobacteria against solar radiation. Journal of Plant Physiology 145, 750-755.
58. Downing, J. E. G., Djamgoz, M. B. A. & Bowmaker, J. K. (1986). Photoreceptors of cyprinid fish: Morphological and spectral characteristics. Journal of Comparative Physiology 159, 859-868.
59. Duchon, J., Borovansky, J. & Hach, P. (1973). Chemical composition of ten kinds of various melanosomes. In V. J. McGovern and P. Russell. Mechanisms in Pigmentation. Pigment Cell 1, 165-170.
60. Dunford, R., Salinaro, A., Cai, L., Serpone, N., Horikoshi, S., Hidaka, H. & Knowland, J. (1997). Chemical oxidation and DNA damage catalyzed by inorganic sunscreen. FEBS Letters, 418, 87-90.
61. Dunlap, W. C. & Chalker, B. E. (1986). Identification and quantitation of near-UV absorbing compounds (S-320) in a hermatypic scleractinian. Coral Reefs 5, 1-5.
62. Dunlap, W. C., Williams, D. Mc B., Chalker, B. E. & Banaszak, A. T. (1989). Biochemical photoadaptation in vision: Uv-absorbing pigments in fish eye tissues. Comparative Biochemistry and Physiology 93 B, 601-607.
63. Dunlap, W. C. & Shick, J. M. (1998). Ultraviolet radiation absorbing mycosporine-like amino acids in coral reef organisms: A biochemical and environmental perspective. Journal of Phycology 34, 418-430.
64. Dunlap, W. C., Chalker, B. E. & Oliver, J. K. (1986). Bathymetric adaptations of reef-building corals at Davis Reef, Great Barrier Reef, Australia. III. UV-B absorbing compounds. Journal of Experimental Marine Biology and Ecology 104, 239-248.
65. Dunlap, W. C. & Yamamoto, Y. (1995). Small-molecule antioxidants in marine organisms: Antioxidant activity of mycosporine-glycine. Comparative Biochemistry and Physiology 112, 105-114.
66. Ehling-Shulz, M., Bilger, W. & Scherer, S. (1997). UV-B induced synthesis of photoprotective pigments and extracellular polysaccharides in the terrestrial cyanobacterium Nostoc commune. Journal of Bacteriology 179, 1940-1945.
67. Ellis, J. & Schramm, D. N. (1995). Could a nearby supernova explosion have caused a mass extinction Proceedings of the National Academy of Sciences, USA 92, 235-238.
68. Everett, M. A., Yeagers, E., Sayre, R. M. & Olson, R. L. (1966). Penetration of epidermis by ultraviolet rays. Photochemistry and Photobiology 5, 533-542.
69. Favre-Bonvin, J., Arpin, N. & Brevard, C. (1976). Structure de la mycosporine. Canadian Journal of Chemistry 54, 1105-1113.
70. Favre-Bonvin, J., Bernillon, J., Salin, N. & Arpin, N. (1987). Biosynthesis of mycosporines: Mycosporine-glutaminol in Trichotecium roseum. Phytochemistry 26, 2509-2514.
71. Flint, S. D. & Caldwell, M. M. (1983). Influence of floral properties on the ultraviolet radiation environment of pollen. American Journal of Botany 70, 1416-1419.
72. Flint, S. D., Jordan, P. W. & Caldwell, M. M. (1985). Plant protective response to enhanced UV-B radiation under field conditions: Leaf optical properties and photosynthesis. Photochemistry and Photobiology 41, 95-99.
73. 2, in glasses of ethanol and ethanediol/water at 77-125K. Journal of the Chemical Society Faraday Transactions 2, 77, 1521-1534.
74. Francois, L. M. & Gerard, J.-C. (1988). Ozone, climate and biospheric environment in the ancient oxygen-poor atmosphere. Planetary and Space Science 36, 1391-1414.
75. Friedmann, P. S. & Gilchrest, B. A. (1987). Ultraviolet radiation directly induces pigment production by cultures human melanocytes. Journal of Cellular Physiology 133, 88-94.
76. Garcia-Pichel, F. (1994). A model for internal self-shading in planktonic organisms and its implications for the usefulness of ultraviolet sunscreens. Limnology and Oceanography 39, 1704-1717.
77. Garcia-Pichel, F. (1998). Solar ultraviolet and evolutionary history of cyanobacteria. Origins of Life and Evolution of the Biosphere 28, 321-347.
78. Garcia-Pichel, F. & Bebout, B. M. (1996). Penetration of ultraviolet radiation into shallow water sediments: High exposure for photosynthetic communities. Marine Ecology Progress Series 131, 257-262.
79. Garcia-Pichel, F. & Belnap, J. (1996). Microenvironments and microscale productivity of cyanobacterial desert crusts. Journal of Phycology 32, 774-782.
80. Garcia-Pichel, F. & Castenholz, R. W. (1991). Characterization and biological implications of scytonemin, a cyanobacterial sheath pigment. Journal of Phycology 27, 395-409.
81. Garcia-Pichel, F. & Castenholz, R. W. (1993). Occurrence of UV-absorbing, mycosporine-like compounds among cyanobacterial isolates and an estimate of their screening capacity. Applied and Environmental Microbiology 59, 163-169.
82. Garcia-Pichel, F., Sherry, N. D. & Castenholz, R. W. (1992). Evidence for an ultraviolet sunscreen role of the extracellular pigment scytonemin in the terrestrial cyano-bacterium Chlorogloeopsis sp. Photochemistry and Photobiology 56, 17-23.
83. Garcia-Pichel, F., Wingard, C. & Castenholz, R. W. (1993). Evidence regarding the UV sunscreen role of a mycosporine-like compound in the cyanobacterium Gloeocapsa sp. Applied and Environmental Microbiology 59, 170-176.
84. Geis, P. A., & Szaniszlo, P. J. (1984). Carotenoid pigments of the dematiaceous fungus Wangiella dermatitidus. Mycologia 76, 268-273.
85. Gilchrest, B. A., Park, H.-Y., Eller, M. S. & Yaar, M. (1996). Mechanisms of ultraviolet-induced pigmentation. Photochemistry and Photobiology 63, 1-10.
86. Gillam, A. E. (1970). Introduction to Electronic Absorption Spectroscopy in Organic Chemistry (eds E. S. Stern and C. J. Timmons), pp. 277. Edward Arnold, London.
87. Gleason, D. F. (1993). Differential effects of ultraviolet radiation on green and brown morphs of the Caribbean coral Porites astreoides. Limnology and Oceanography 38, 1452-1463.
88. Grant, P. T., Middleton, C., Plack, P. A. & Tomson, R. H. (1985). The isolation of four aminocyclohexenimines (mycosporines) and a structurally related derivative of cyclohexane-1,3-dione (gadusol) from the brine shrimp. Artemia Comparative Biochemistry Physiology 80 B, 755-759.
89. Grant, P. T. & Plack, P. A. (1980). Gadusol, a metabolite from fish eggs. Tetrahedron Letters 21, 4043-4044.
90. Groth, W. & Weyssenhoff, H. V. (1960). Photochemical formation of organic compounds from mixtures of simple gases. Planetary and Space Science 2, 79.
91. 13CNMR spectroscopy of sporopollenins from different plant taxa. Plant Physiology 86, 134-136.
92. Häder, D.-P. & Worrest, R. C. (1991). Effects of enhanced solar ultraviolet radiation on aquatic ecosystems. Photochemistry and Photobiology 53, 717-725.
93. Hadwiger, L. A. & Schwochau, M. E. (1971). Ultraviolet light induced formation of pisatin and phenylalanine ammonia-lyase. Plant Physiology 47, 588-590.
94. Hanawalt, P. C., Cooper, P. K., Ganesan, A. K. & Smith, C. A. (1979). DNA repair in bacteria and mammalian cells. Annual Review of Biochemistry 48, 783-836.
95. Harborne, J. B. (1964). Biochemistry of Phenolic Compounds. Academic Press, London.
96. Harm, W. (1980). Biological Effects of Ultraviolet Radiation. IUPAB Biophysics Series I. Cambridge University Press, Cambridge.
97. Harosi, F. I. & Hashimoto, Y. (1983). Ultraviolet sensitivity in goldfish: An independent UV retinal mechanism. Vision Research 25, 11-20.
98. Hawksworth, D. L. & Hill, D. J. (1984). The Lichen Forming Fungi. Blackie, Glasgow.
99. Haxo, F. T., Lewin, R. A., Lee, K. W. & Li, M.-R. (1987). Fine structure and pigments of Oscillatoria (Trichodesmium) aff. thiebautii (Cyanophyta) in culture. Phycologia 26, 443-456.
100. Holbrook, K. A. & Wulff, K. (1987). The structure and development of skin. In Dermatology in General Medicine (eds T. B. Fitzpatrick, A. Z. Eisen, K. Wulff and K. F. Austen), pp. 93-131. McGraw-Hill, New York.
101. Horodyski, R. J. & Knauth, L. P. (1994). Life on land in the Precambrian. Science 263, 494-498.
102. Ito, S. & Hirata, T. (1977). Isolation and structure of a mycosporine from the zoanthid Palythoa tuberculosa. Tetrahedron Letters 28, 2429-2430.
103. Jagger, J. (1985). Solar-UV Actions on Living Cells. Praeger Scientific, New York.
104. Johnston, D. A. (1980). Volcanic contribution of chlorine to the stratosphere: More significant to ozone that previously estimated Science 209, 494-493.
105. Jorgensen, R. (1993). The origin of land plants: A union of algae and fungi advanced by flavonoids. Biosystems 31, 193-207.
106. 2. Chemistry in Britain, 503-505.
107. Kaidbey, K. H., Poh Agin, P., Sayre, R. M. & Kligman, A. M. (1979). Photoprotection by melanin - a comparison of black and caucasian skin. Journal of the American Academy of Dermatology 1, 249-260.
108. Karabourniotis, G., Papadopoulos, K., Papamarkou, M. & Manetas, Y. (1992). Ultraviolet-B radiation absorbing capacity of leaf hairs. Physiologia Plantarum 86, 414-418.
109. Karentz, D., Bosch, I. & Dunlap, W. C. (1992). Distribution of UV-absorbing compounds in the antarctic limpet, Nacella concinna. Antarctic Journal 27, 121-122.
110. Karentz, D., Cleaver, J. E. & Mitchell, D. L. (1991a). Cell survival characteristics and molecular responses of Antarctic phytoplankton to ultraviolet-B radiation. Journal of Phycology 27, 326-341.
111. Karentz, D., Dunlap, W. C. & Bosch, I. (1997). Temporal and spatial occurrence of UV-absorbing mycosporine-like amino acids in tissues of the antarctic sea urchin Sterechinus neumayeri during springtime ozone-depletion. Marine Biology 129, 343-353.
112. Karentz, D., McEuen, F. S., Land, M. C. & Dunlap, W. C. (1991b). Survey of mycosporine-like amino acids in Antarctic marine organisms: Potential protection from ultraviolet exposure. Marine Biology 108, 157-166.
113. Kasting, J. F. (1987). Theoretical constraints on oxygen and carbon dioxide concentrations in the precambrian atmosphere. Precambrian Research 34, 205.
114. Kasting, J. F. (1993). Earth's early atmosphere. Science 259, 920-926.
115. Kasting, J. F. & Donahue, T. M. (1980). The evolution of atmospheric ozone. Journal of Geophysical Research 85, 3255.
116. Kasting, J. F., Zahnle, K. J., Pinto, J. P. & Young, A. T. (1989). Sulfur, ultraviolet radiation and the early evolution of life. Origins of Life and Evolution of the Biosphere 19, 95-108.
117. Kligman, L. H., Akin, F. J. & Kligman, A. M. (1980). Sunscreens prevent ultraviolet photocarcinogenesis. Journal of the American Academy of Dermatology 3, 30-35.
118. Knoll, A. H. (1982). Microfossils from late precambrian Draken conglomerate. Ny. Friesland, Svalbard. Journal of Palaeontology 56, 755-790.
119. Knowland, J., McKenzie, E. A., McHugh, P. J. & Cridland, N. A. (1993). Sunlight-induced mutagenicity of a common sunscreen ingredient. FEBS Letters 324, 309-313.
120. Koes, R. E., Quattrocchio, F. & Mol, J. N. M. (1994). The flavonoid biosynthetic pathway in plants: Function and evolution. Bioessays 16, 123-132.
121. Kolb, V. M., Dworkin, J. P. & Miller, S. L. (1994). Alternative bases in the RNA world: The prebiotic synthesis of urazole and its ribosides. Journal of Molecular Evolution 38, 549-557.
122. Kollias, N., Sayre, R. M., Zeise, L. & Chedekel C., (1991). Photoprotection by melanin. Journal of Photochemistry and Photobiology 9, 135-160.
123. Krinsky, N. I. (1971). Function (carotenoids). In Carotenoids (ed. O. Isler), pp. 669-716. Halsted Press, John-Wiley, New York.
124. Krinsky, N. I. (1979). Carotenoid protection against oxidation. Pure and Applied Chemistry 51, 649-660.
125. Kubota, Y., Shuin, T., Kawasaki, C., Hosaka, H., Kitamura, H., Cai, R., Sakai, H., Hashimoto, K. & Fujishima, A. (1994). Photokilling of T-24 human bladder cancer cells with titanium dioxide. British Journal of Cancer 70, 1107-1111.
126. Larson, R. A. (1988). The antioxidants of higher plants. Photochemistry 27, 969-978.
127. Lazcano, A. (1994). The RNA world, its predecessors, and its descendants. In Early Life on Earth (ed. S. Bengtson), pp. 70-80. Columbia University Press, New York.
128. Leach, C. M. (1965). Ultraviolet absorbing substances associated with light-induced sporulation in fungi. Canadian Journal of Botany 43, 185-200.
129. Leavitt, P. R., Vinebrook, R. D., Donald, D. B., Smol, J. P. & Schindler, D. W. (1997). Past ultraviolet radiation environments in lakes derived from fossil pigments. Nature 388, 457-459.
130. Levin, D. A. (1976). Alkaloid-bearing plants: An ecogeographic perspective. American Naturalist 110, 261-284.
131. Llewellyn, C. A. & Mantoura, R. F. C. (1997). A UV-absorbing compound in HPLC pigment chromatograms obtained from Icelandic basin phytoplankton. Marine Ecology Progress Series 158, 283-287.
132. Lowry, B., Lee, D. & Hebant, C. (1980). The origin of land plants: A new look at an old problem. Taxon 29, 183-197.
133. Lyndon, J., Teramura, A. H. & Coffman, C. B. (1987). UV-B radiation effects on photosynthesis, growth and cannabinoid production of two Cannabis sativa chemotypes. Photochemistry and Photobiology 46, 201-206.
134. Makino, C. L., Dodd, R. L., Rohlich, P. & Baylor, D. A. (1995). Salamander UV-sensitive cones utilize more than one visual pigment. Biophysical Journal 68, A19.
135. Marchant, H. J., Davidson, A. T. & Kelly, G. J. (1991). UV-B protecting compounds in the marine alga Phaeocystis pouchetii from Antarctica. Marine Biology 109, 391-395.
136. Margulis, L., Walker, J. C. G. & Rambler, M. (1976). Reassessment of roles of oxygen and ultraviolet light in Precambrian evolution. Nature 264, 620-624.
137. Markham, K. R. (1988). Distribution of flavonoids in the lower plants and its evolutionary significance. In The Flavonoids, Advances in Research since 1980 (ed. J. B. Harborne), pp. 427-468. Academic Press, New York.
138. Markham, K. R. (1989). Flavones, Flavonols and their glycosides. In Methods in Plant Biochemistry, pp. 197-235. Academic Press, London.
139. Martincigh, B. S., Allen, J. M. & Allen, S. K. (1997). Sunscreens: The molecules and their photoreactivity. In Sunscreen Photobiology (ed. F. P. Gasparro), pp. 11-45, Springer-Verlag, New York.
140. McClure, J. W. (1975). Physiology and function of flavonoids. In The Flavonoids (eds J. B. Harborne, T. J. Marby and H. Mabry) Part 2, pp. 970-1055. Academic Press, New York.
141. McHugh, P. J. & Knowland, J. (1998). Characterisation of DNA damage inflicted by free radicals from a mutagenic sunscreen ingredient and its location using an in vitro genetic reversion assay. Photochemistry and Photobiology 66, 276-281.
142. Menon, I. A., Persad, S., Haberman, H. F. & Kurlan, C. J. (1983). A comparative study of the physical and chemical properties of melanins isolated from human red and black hair. Journal of Investigative Dermatology 80, 202-206.
143. Meuter-Gerhards, A., Schwerdtfeger, C. & Steuernagel, S. (1995). Studies on sporopollenin structure during pollen development. Zeitschrift fuer Naturforschung 50, 487-492.
144. Meyer-Rochow, V. B. (1991). Differences in ultraviolet wing patterns in the New Zealand lucaenid butterflies, Lucaena salustius, Lucaena rauparaha, and Lucaena feredayi as a likely isolating mechanism. Journal of the Royal Society of New Zealand 21, 169-178.
145. Mohle, B. & Wellmann, E. (1982). Induction of phenyl-propanoid compounds by UV-B irradiation in roots of seedlings and cell cultures from Dill (Anethum gaveolens L.) Plant Cell Reports 1, 183-185.
146. Moorthy, P. & Kathiresan, K. (1997). Photosynthetic pigments in tropical mangroves: Impacts of seasonal flux of UV-B radiation and other environmental attributes. Botanica Marina 40, 341-349.
147. Mulkidjanian, A. Y. & Junge, W. (1997). On the origin of photosynthesis as inferred from sequence analysis. Photosynthesis Research 51, 27-42.
148. Mulroy, T. W. (1979). Spectral properties of heavily glaucous and non-glaucous leaves of a succulent rosette-plant. Oecologia 38, 349-357.
149. Muntz, W. R. A. (1973). Yellow filters and the absorption of light by the visual pigments of some Amazonian fishes. Vision Research 13, 2235-2254.
150. Nägeli, C. & Schwenderer, S. (1877). Das Mikroskop. W. Englemann Verlag, Leipzig.
151. Nakamura, H., Kobayashi, J. & Hirata, Y. (1981). Isolation and structure of a 330 nm UV-absorbing substance, asterina 330 from the starfish, Asterina pectinifera. Chemical Letters 1981, 1413-1414.
152. Nakamura, H., Kobayashi, J. & Hirata, Y. (1982). Separation of mycosporine-like amino acids in marine organisms using reversed phased high performance liquid chromatography. Journal of Chromatography 250, 113-118.
153. Natow, A. J. (1986). Sunscreens. Cutis 38, 157-158.
154. Nicolet, M. (1989). Solar spectral irradiances and their diversity between 120 and 900 nm. Planetary and Space Science 37, 1249-1289.
155. Nienow, J. A. & Friedmann, E. I. (1993). Terrestrial litho-phytic (rock) communities. In Antarctic Microbiology (ed. E. I. Friedmann), pp. 343-412. Wiley-Liss, Inc, New York.
156. Nienow, J. A., McKay, C. P. & Friedmann, E. I. (1988). The cryptoendolithic microbial environment in the Ross Desert of Antarctica: Light in the photosynthetically active region. Microbial Ecology 16, 271-289.
157. Niklas, K. J. (1976). Morphological and chemical examinations of Courviosiella ctenomorpha gen. and sp. nov.: A siphonous alga from the Upper Devonian, West Virginia, U.S.A. Review of Palaeobotany and Palynology 21, 187-203.
158. Noorudeen, A. M. & Kulandaivelu, G. (1982). On the possible site of inhibition of photosynthetic electron transport by ultraviolet-B (UV-B) radiation. Physiologia Plantarum 55, 161-166.
159. Nordlund, J. J. & Boissy, R. (1998). The Pigmentary System. Oxford University Press, Oxford.
160. Ogura, N. & Hanya, T. (1966). Nature of ultra-violet absorption of sea water. Nature 212, 758.
161. Olsen, J. M. & Pierson, B. K. (1986). Photosynthesis 3.5 thousand million years ago. Photosynthesis Research 9, 251-259.
162. Oren, A. (1997). Mycosporine-like amino acids as osmotic solutes in a community of halophilic cyanobacteria. Geomicrobiology Journal 14, 231-240.
163. Patel, N. P., Highton, A. & Moy, R. L. (1992). Properties of topical sunscreen formulations. Journal of Dermatology and Surgical Oncology 18, 316-320.
164. Pentecost, A. (1993). Field relationships between scytonemin density, growth and irradiance in cyanobacteria occurring in low illumination regimes. Microbial Ecology 26, 101-110.
165. Pierson, B. K., Mitchell, H. K. & Ruff-Roberts, A. L. (1993). Chloroflexus aurantiacus and ultraviolet radiation: Implications for archean shallow-water stromatolites. Origins of Life and Evolution of the Biosphere 23, 243-260.
166. Plack, P. A., Fraser, N. W., Grant, P. T., Middleton, C., Mitchell, A. I. & Thomson, R. H. (1981). Gadusol, an enolic derivative of cyclohexane-1,3-dione present in the roes of cod and other marine fish. Biochemical Journal 199, 741-747.
167. Prezelin, B. B., Boucher, N. P. & Schofield, O. (1994). Evaluation of field studies of UV-B radiation effects on Antarctic marine primary productivity. In Stratospheric Ozone Depletion)UV-B Radiation in the Biosphere. Springer-Verlag, New York.
168. Prota, G. (1988). Progress in the chemistry of melanins and related metabolites. Medical Research Reviews 8, 525-556.
169. Proteau, P. J., Gerwick, W. H., Garcia-Pichel, F. & Castenholz, R. (1993). The structure of scytonemin: An ultraviolet sunscreen pigment from the sheaths of cyanobacteria. Experientia 49, 825-829.
170. Rambler, M. B. & Margulis, L. (1980). Bacterial resistance to ultraviolet irradiation under anaerobiosis: Implications for pre-phanerozoic evolution. Science 210, 638-640.
171. Rampino, M. R., Self, S. & Stothers, R. B. (1988). Volcanic winters. Annual Reviews of Earth and Planetary Sciences 16, 73-99.
172. Ravishankar, J. P., Muruganandam, V. & Suryanarayanan, T. S. (1995). Isolation and characterization of melanin from a marine fungus. Botanica Marina 38, 413-416.
173. Renger, G., Volker, M., Eckerts, H. J., Fromme, R., Hohm-Veit, S. & Graber, P. (1989). On the mechanism of photosystem II deterioration by UV-B radiation. Photochemistry and Photobiology 49, 97-105.
174. Robberecht, R. & Caldwell, M. M. (1978). Leaf epidermal transmittance of ultraviolet radiation and its implications for plant sensitivity to ultraviolet-radiation induced injury. Oecologia 32, 277-287.
175. Robberecht, R., Caldwell, M. M. & Billings, W. D. (1980). Leaf ultraviolet optical properties along a latitudinal gradient in the arctic-alpine life zone. Ecology 61, 612-619.
176. Roscher, N. M., Lindemannm, M. K. O., Kong, S. B., Cho, C. G. & Jiang, P. (1994). Photodecomposition of several compounds commonly used as sunscreen agents. Journal of Photochemistry and Photobiology A: Chem. 80, 417-421.
177. Rothschild, L. J. (1990). Earth analogs for martian life. Microbes in evaporites. A new model system for life on Mars. Icarus 88, 246-260.
178. Ruderman, M. A. (1974). Possible consequences of nearby supernova explosions for atmospheric ozone and terrestrial life. Science 184, 1079.
179. Sagan, C. (1973). Ultraviolet radiation selection pressure on the earliest organisms. Journal of Theoretical Biology 39, 195-200.
180. Sagan, C. & Khare, B. N. (1971). Long wave UV photo-products of amino acids on the primitive Earth. Science 173, 417.
181. Salisbury, F. B. & Ross, C. W. (1992). Plant Physiology. Wadsworth, Belmont.
182. Scelfo, G. M. (1988). Ultraviolet-B absorbing compounds in Anthopleura elegantissima. American Zoologist 28, 105.
183. Scherer, S., Chen, T. W. & Boger, P. (1988). A new UV-A/B protecting pigment in the terrestrial cyanobacteria Nostoc commune. Plant Physiology 88, 1055-1057.
184. Schopf, J. W. & Klein, C. (1992). The Proterozoic Biosphere: A Multidisciplinary Study. Cambridge University Press, Cambridge.
185. Schwack, W. & Rudolph, T. (1995). Photochemistry of dibenzoyl methane UV-A filters, Part 1. Journal of Photochemistry and Photobiology B: Biol. 28, 229-234.
186. Scotto, J. & Fraummeni, J. F. (1982). Skin cancer (other than melanoma). In Cancer Epidemiology and Prevention (ed. D. Scottenfeld), ch. 60, 996-1011. Saunders, Philadelphia.
187. Sekikawa, I., Kubota, C., Hiraoki, T. & Tsujino, I. (1986). Isolation and structure of a 357 nm UV-absorbing substance, usujirene, from the red alga Palmaria palmata (L.) O. Kuntze. Japanese Journal of Phycology 34, 185-188.
188. Serpone, N. (1996). Applied heterogeneous photocatalysis with semiconductor materials-photocatalyzed detoxification of wastewaters and air. Kirk-Othmer Encyclopedia of Chemical Technology, 18, 820-837.Wiley-Interscience, New York.
189. Setlow, R. B., Grist, E., Thompson, K. & Woodhead, A. D. (1993). Wavelengths effective in induction of malignant melanoma. Proceedings of the National Academy of Sciences, USA 90, 6666-6670.
190. Sharma, N., Hepburn, D. & Fitt, P. S. (1984). Photo-reactivation in pigmented and non-pigmented extreme halophiles. Biochimica et Biophysica Acta 799, 135-142.
191. Shibata, K. (1969). Pigments and a UV-absorbing substance in corals and a blue-green alga living in the Great Barrier Reef. Plant and Cell Physiology 10, 741-747.
192. Shick, J. M., Dunlap, W. C., Chalker, B. E., Banaszak, A. T. & Rosenzweig, T. K. (1992). Survey of ultraviolet radiation-absorbing mycosporine-like amino acids in organs of carol reef holothuriods. Marine Ecology Progress Series 90, 139-148.
193. Shick, J. M., Dunlap, W. C. & Larson, R. M. (1994). Coral reef holothuriods (Echinodermata) accumulate UV-photoprotectants from their diet. American Zoologist 33, 62.
194. Shick, J. M., Lesser, M. P., Dunlap, W. C., Stochaj, W. R., Chalker, B. E. & Wu Won, J. (1995). Depth-dependent responses to solar ultraviolet radiation and oxidative stress in the zooanthellate coral Acropora microphthalma. Marine Biology 122, 41-51.
195. Shick, J. M., Lesser, M. P. & Stochaj, W. R. (1991). Ultraviolet radiation and photooxidative stress in zooanthellate Anthozoa: The sea anemone Phyllodiscus semoni and the octocoral Clavularia sp. Symbiosis 10, 145-173.
196. Shimazaki, K., Igarashi, T. & Kondo, N. (1988). Protection of the epidermis of photosynthesis against UV-C radiation estimated by chlorophyll a florescence. Physiologia Plantarum 74, 34-38.
197. Sinha, R. P., Singh, N., Kumar, A., Kumar, H. D., Häder, M. & Häder, D. P. (1996). Effects of UV irradiation on certain physiological and biochemical processes in cyanobacteria. Journal of Photochemistry and Photobiology 32, 107-113.
198. Sivalingam, P. M., Ikawa, T. & Nisizawa, K. (1974a). Possible physiological roles of a substance showing characteristic UV-absorbing patterns in some marine algae. Plant and Cell Physiology 15, 583-586.
199. Sivalingam, P. M., Ikawa, T., Yokohama, Y. & Nisizawa, K. (1974b). Distribution of a 334 UV-absorbing-substance in algae, with special regard of its possible physiological roles. Botanica Marina 17, 23-29.
200. Smith, R. C., Baker, K. S., Holm-Hansen, O. & Olsen, R. (1980). Photoinhibition of photosynthesis in natural waters. Photochemistry and Photobiology 31, 585-592.
201. Smith, R. C., Prezelin, B. B., Baker, K. S., Bidigare, R. R., Boucher, N. P., Coley, T., Karentz, D., MacIntyre, S., Matlick, H. A., Menzies, D., Ondrusek, M., Wan, Z. & Waters, K. J. (1992). Ozone depletion: Ultraviolet radiation and phytoplankton biology in Antarctic waters. Science 255, 952-959.
202. Solhaug, K. A. & Gauslaa, Y. (1996). Parietin, a photoprotective secondary product of the lichen Xanthoria parietina. Oecologia 108, 412-418.
203. Spiegel-Adolf, M. (1937). Studies on melanins. I. Photosynthetic melanins. Biochem J. 31, 1303-1310.
204. Stafford, H. A. (1991). Flavonoid evolution: An enzymatic approach. Plant Physiology 96, 680-685.
205. Stochaj, W. R., Dunlap, W. C. & Shick, J. M. (1994). Two new UV-absorbing mycosporine-like amino acids from the sea anemone Anthopleura elegantissima and the effects of zooxanthellae and spectral irradiance on chemical composition and content. Marine Biology 118, 149-156.
206. Strack, D. & Wray, V. (1989). Anthocyanins. In Methods in Plant Biochemistry, pp. 325-356. Academic Press, London.
207. Sutherland, B. M. (1981). Photoreactivation. Bioscience 31, 439-444.
208. Sutherland, B. M. (1982). p-Aminobenzoic acid-sunlamp sensitization of pyrimidine dimer formation and transformation in human cells. Photochemistry and Photobiology 36, 95-97.
209. Sutherland, J. C. & Griffin, K. P. (1984). p-aminobenzoic acid can sensitize the formation of pyrimidine dimers in DNA: Direct chemical evidence. Photochemistry and Photobiology 40, 391-394.
210. Takano, S., Nakanishi, A., Uemura, D. & Hirata, Y. (1979). Isolation and structure of a 334 nm UV-absorbing substance, porphyra-334 from the red alga Porphyra tenera Kjellman. Chemical Letters 1979, 419-420.
211. Takano, S., Uemura, D. & Hirata, Y. (1978a). Isolation and structure of a new amino acid, palythine, from the zoanthid Palythoa tuberculosa. Tetrahedron Letters 26, 2299-2300.
212. Takano, S., Uemura, D. & Hirata, Y. (1978b). Isolation and structure of two amino acids, palythinol and palythene, from the zoanthid Palythoa tuberculosa. Tetrahedron Letters 49, 4909-4912.
213. Tang, E. P. Y., Tremblay, R. & Vincent, W. F. (1997). Cyanobacterial dominance of polar freshwater ecosystems: Are high latitude mat-formers adapted to low temperature Journal of Phycology 33, 171-181.
214. Tevini, M. (1993). UV-B Radiation and Ozone Depletion. Effects on Humans, Animals, Plants, Micro-organisms and Materials. Lewis Publishers, Boca Raton, Florida.
215. Tevini, M., Braun, J. & Fieser, G. (1991). The protective function of the epidermal layer of rye seedlings against ultraviolet-B radiation. Photochemistry and Photobiology 53, 329-333.
216. Tevini, M. & Iwanzik, W. (1983). Inhibition of photosynthetic activity by UV-B radiation in radish seedlings. Physiologia Plantarum 58, 395-400.
217. Tevini, M. & Teramura, A. H. (1989). UV-B effects on terrestrial plants. Photochemistry and Photobiology 50, 479-487.
218. Thorsett, S. E. (1995). Terrestrial implications of cosmological gamma-ray burst models. The Astrophysical Journal 444, L53.
219. Toon, O. B., Zahnle, K., Morrison, D., Turco, R. P. & Covey, C. (1997). Environmental perturbations caused by the impacts of asteroids and comets. Reviews of Geophysics 35, 41-78.
220. Trent, J. D., Kagawa, H. K., Yaqi, T., Olle, E. & Zaluzec, N. J. (1997). Chaperonin filaments: The archeal cytoskeleton Proceedings of the National Academy of Sciences, USA 94, 5383-5388.
221. Tsujino, I., Yabe, K., Sekikawa, I. & Hamanaka, N. (1978). Isolation and structure of a mycosporine from the red alga Chondrus yendroi. Tetrahedron Letters 16, 1401-1402.
222. Tsujino, I., Yabe, K. & Sekikawa, I. (1980). Isolation and structure of a new amino acid, shinorine, from the red algae Chondrus yendoi Yamada et Mikami. Botanica Marina 23, 65-68.
223. Turco, R. P., Toon, O. B., Park, C., Whitten, R. C. & Pollack, J. B. (1982). An analysis of the physical, chemical, optical and historical impacts of the 1908 Tunguska meteor fall. Icarus 50, 1-52.
224. Van der Leun, J. C. & De Gruijl, F. R. (1993). Influences of ozone depletion on human and animal health. In UV-B radiation and ozone depletion. Effects on humans, animals, plants, micro-organisms and materials (ed. M. Tevini). Lewis Publishers, Boca Raton, FL.
225. Vincent, W. F. & Quesada, A. (1994). Ultraviolet radiation effects on cyanobacteria: Implications for Antarctic microbial ecosystems. In Ultraviolet Radiation in Antarctica: Measurements and biological effects (eds. C. S. Weiler and P. A. Penhale). Antarctic Research Series 62, 111-124.
226. Vincent, W. F. & Roy, S. (1993). Solar ultraviolet-B radiation and aquatic primary production: Damage, protection and recovery. Environmental Reviews 1, 1-12.
227. Vogelmann, A. M., Ackerman, T. P. & Turco, R. P. (1992). Enhancements of biologically effective ultraviolet radiation following volcanic eruptions. Nature 359, 47-49.
228. Vogt, T., Gulz, P. & Reznik, H. (1991). UV Radiation dependent flavonoid accumulation of Cistus laurifolius L. Z. Naturforsch 46, 37-42.
229. Wamer, W. G., Yin, J.-J., & Wei, R. R. (1997). Oxidative damage to nucleic acids photosensitized by titanium dioxide. Free Radical Biology and Medicine 23, 851-858.
230. Wangberg, S., Persson, A. & Karlson, B. (1997). Effects of UV-B radiation on synthesis of mycosporine-like amino acid and growth in Heterocapsa triquetra (Dinophyceae). Journal of Photochemistry and Photobiology 37, 141-146.
231. Wayne, R. P. (1988). Principles and Applications of Photochemistry, Oxford University Press, Oxford.
232. Weissenbock, G., Plesser, A. & Trinks, K. (1976). Flavonoidgehalt und enzymaktivitaten isolierter haferchloroplasten (Avena sativa L.) Ber. Dtsch. Bot. Ges. 89, 457-472.
233. Wellmann, E. (1975). UV dose-dependent induction of enzymes related to flavonoid biosynthesis in cell suspension cultures of parsley. FEBS Letters 51, 105-107.
234. Wellman, E., Hrazdina, G. & Grisebach, H. (1976). Induction of anthocyanin formation and of enzymes related to its biosynthesis by UV light in cell cultures of Haplopappus gracilis. Phytochemistry 15, 913-915.
235. 13C/MAS NMR spectroscopic investigations on sporopollenin of different systematic origins. Z. Naturforsch 48, 697-701.
236. Wulf, H. C., Poulsen, T., Brodthagen, H. & Hou-Jensen, K. (1982). Sunscreens for delay of ultraviolet induction of skin tumours. Journal of the American Academy of Dermatology 7, 194-202.
237. Xiong, F., Komenda, J., Kopecky, J. & Nedbal, L. (1997). Strategies of ultraviolet-B protection in microscopic algae. Physiologia Plantarum 100, 378-388.
238. Xiong, F., Lederer, F., Lukavsky, J. & Nedbal, L. (1996). Screening of freshwater algae (Chlorophyta, Chromophyta) for ultraviolet-B sensitivity of the photosynthetic apparatus. Journal of Plant Physiology 148, 42-48.
239. Young, H. & Patterson, V. J. (1982). A UV photoprotective compound from Glomerella cingulata- a mycosporine. Phytochemistry 21, 1075-1077.
240. Zangerl, A. R. & Berenbaum, M. R. (1987). Furanocoumarins in wild parsnip: Effect of photosynthetically active radiation, ultraviolet light and nutrients. Ecology 68, 516-520.
241. Zigman, S. (1987). Biochemical adaptation in vision. Photochemistry and Photobiology 45s, 35.
242. Zigman, S., Paxia, T. & Waldron, W. (1985). Properties and functions of near UV absorbing pigments in marine animal lenses. Biological Bulletin 169, 564.