• 1
    Vincent, W. F. and P. J. Neale (2000) Mechanisms of UV damage to aquatic organisms. In The Effects of UV Radiation in the Marine Environment (Edited by S. de Mora, S. Demers and M. Vernet), pp. 149176. Cambridge University Press, Cambridge.
  • 2
    Karentz, D. (2001) Chemical defenses of marine organisms against solar radiation exposure: UV-absorbing mycosporine-like amino acids and scyotnemin. In Marine Chemical Ecology (Edited by J. B. McClintock and B. J. Baker), pp. 481486. CRC Press, Birmingham.
  • 3
    Hoffman, J., L. J. Hansen and T. Klinger (2003) Interactions between UV radiation and temperature limit inferences from single-factor experiments. J. Phycol. 39, 268272.
  • 4
    Bischof, K., I. Gómez, M. Molis, D. Hanelt, U. Karsten, U. Lüder, M. Y. Roleda, K. Zacher and C. Wiencke (2006) UV radiation shapes seaweed communities. Rev. Environ. Sci. Biotechnol. 5, 141166.
  • 5
    Cie, D. K. and M. S. Edwards (2008) The effects of high radiation on the settlement competency and viability of kelp zoospores. J. Phycol. 44, 495500.
  • 6
    Karsten, U., A. Wulff, M. Y. Roleda, R. Müller, F. S. Steinhoff, J. Fredersdorf and C. Wiencke (2009) Physiological responses of polar benthic algae to ultraviolet radiation. Bot. Mar. 52, 639654.
  • 7
    Bischof, K., D. Hanelt, H. Tüg, U. Karsten, P. E. M. Brouwer and C. Wiencke (1998) Acclimation of brown algal photosynthesis to ultraviolet radiation in Arctic coastal waters (Spitsbergen, Norway). Polar Biol. 20, 388395.
  • 8
    Goes, J. I., N. Handa, S. Taguschi and T. Hama (1994) Effect of UV-B radiation on the fatty acid composition of the marine phytoplankter Tetraselmis sp.: Relationship to cellular pigments. Mar. Ecol. Prog. Ser. 114, 259274.
  • 9
    Leu, E., S. Falk-Petersen and D. O. Hessen (2007) Ultraviolet radiation negatively affects growth but not food quality of arctic diatoms. Limnol. Oceanogr. 52, 787797.
  • 10
    Zacher, K. (2007) Effects of UV-Radiation on Antarctic Benthic Algae with Emphasis on Early Successsional Stages and Communities. Ph.D. thesis, University of Bremen.
  • 11
    Roleda, M. Y., C. Wiencke, D. Hanelt and K. Bischof (2007) Sensitivity of the early life stages of macroalgae from the northern hemisphere to ultraviolet radiation. Photochem. Photobiol. 83, 851862.
  • 12
    Wang, K. S. and T. J. Chai (1994) Reduction of omega-3 fatty acids by UV-B radiation in microalgae. J. Appl. Phycol. 6, 415421.
  • 13
    Hessen, D. O., H. J. De Lange and E. van Donk (1997) UV-induced changes in phytoplankton cells and its effects on grazers. Freshwater Biol. 38, 513524.
  • 14
    Wiencke, C., I. Gómez, H. Pakker, A. Flores-Moya, M. Altamirano, D. Hanelt, K. Bischof and F. L. Figueroa (2000) Impact of UV-radiation on viability, photosynthetic characteristics and DNA of brown algal zoospores: Implications for depth zonation. Mar. Ecol. Prog. Ser. 197, 217229.
  • 15
    Wiencke, C., B. Vögele, N. A. Kovalttchouk and H. Hop (2004) Species composition and zonation of marine benthic macroalgae at Hansneset in Kongsfjorden, Svalbard. Ber. Polarforsch. Meeresforsch. 492, 5572.
  • 16
    Wiencke, C., M. Y. Roleda, A. Gruber, M. N. Clayton and K. Bischof (2006) Susceptibility of zoospores to UV radiation determines upper depth distribution limit of Arctic kelps: Evidence through field experiments. J. Ecol. 94, 455463.
  • 17
    Roleda, M. Y., C. Wiencke, D. Hanelt, W. H. van de Poll and A. Gruber (2005) Sensitivity of Laminariales zoospores from Helgoland (North Sea) to ultraviolet and photosynthetically active radiation: Implications for depth distribution and seasonal reproduction. Plant Cell Environ. 28, 466479.
  • 18
    Roleda, M. Y. (2006) Effects of Ultraviolet Radiation on Early Life Stages of Cold Temperate an Arctic Macroalgae: Implications for Recruitments and Vertical Depth Distribution. Ph.D. thesis, University of Bremen.
  • 19
    Roleda, M. Y., M. N. Clayton and C. Wiencke (2006a) Screening capacity of UV-absorbing compounds in spores of Arctic Laminariales. J. Exp. Mar. Biol. Ecol. 338, 123133.
  • 20
    Tala, F., K. Veliz, I. Gomez and M. Edding (2007) Early life stages of the South Pacific kelps Lessonia nigrescens and Lessonia trabeculata (Laminariales, Phyeophyceae) show recovery capacity following exposure to UV radiation. Phycologia 46, 467470.
  • 21
    Müller, R., C. Wiencke and K. Bischof (2008) Interactive effects of UV radiation and temperature on microstages of Laminariales (Phaeophyceae) from the Arctic and North Sea. Clim. Res. 37, 203213.
  • 22
    Müller, R., C. Wiencke, K. Bischof and B. Krock (2009) Zoospores of three Laminariales under different UV radiation and temperature conditions: Exceptional spectral absorbance properties and lack of phlorotannin induction. Photochem. Photobiol. 85, 970977.
  • 23
    Frankel, E. N., W. E. Neff and E. Selke (1981) Analysis of autoxidized fats by gas chromatography–mass spectrometry: VII. Volatile thermal decomposition products of pure hydroperoxides from autoxidized and photosensitized oxidized methyl oleate, linoleate and linolenate. Lipids 16, 279285.
  • 24
    Dennis, K. J. and T. Shibamoto (1990) Gas chromatographic analysis of reactive carbonyl compounds formed from lipids upon UV radiation. Lipids 25, 460464.
  • 25
    Guihéneuf, F., V. Mimouni, L. Ulmann and G. Tremblin (2008) Environmental factors affecting growth and omega 3 fatty acid composition in Skeletonema costatum. The influences of irradiance and carbon source. Diatom Res. 28, 93103.
  • 26
    Guihéneuf, F., V. Mimouni, L. Ulmann and G. Tremblin (2009) Combined effects of irradiance levels and carbon source on fatty acid and lipid class composition in the microalga Pavlova lutheri commonly used in mariculture. J. Exp. Mar. Bot. 369, 136143.
  • 27
    Targett, N. M. and T. M. Arnold (1998) Predicting the effects of brown algal phlorotannins on marine herbivores in tropical and temperate oceans. J. Phycol. 34, 195205.
  • 28
    Jormalainen, V., T. Honkanen, R. Koivikko and J. Eränen (2003) Induction of phlorotannin production in a brown alga: Defense or resource dynamics? Oikos 103, 640650.
  • 29
    Crato, E. (1893) Morphologische und mikrochemische Untersuchungen der Physoden. Bot. Ztg. 51, 157194.
  • 30
    Ragan, M. A. (1976) Physodes and the phenolic compounds of brown algae. Composition as significance of physodes in vivo. Bot. Mar. 19, 145154.
  • 31
    Schoenwaelder, M. E. A. and M. N. Clayton (1999) The role of the cytoskeleton in brown algal physode movement. Eur. J. Phycol. 34, 223229.
  • 32
    Schoenwaelder, M. E. A. and M. N. Clayton (1998) Secretion of phenolic substances into the zygote wall and cell plate in embryos of Hormosira and Acrocarpia (Fucales, Phaeophyceae). J. Phycol. 34, 969980.
  • 33
    Lüder, U. H. and M. N. Clayton (2004) Induction of phlorotannins in the brown macroalga Ecklonia radiata (Laminariales, Phaeophyta) in response to simulated herbivory—The first microscopic study. Planta 218, 928937.
  • 34
    Ragan, M. A. and K. W. Glombitza (1986) Phlorotannins, brown algal polyphenols. Prog. Phycol. Res. 4, 129241.
  • 35
    Waterman, P. G. and S. Mole (1994) Analysis of Phenolic Plant Metabolites. Blackwell Scientific Publications, Oxford, Great Britain.
  • 36
    Arnold, T. M. and N. M. Targett (2002) Marine tannins: The importance of a mechanistic framework for predicting ecological roles. J. Chem. Ecol. 28, 19191934.
  • 37
    Steinhoff, F. S., M. Graeve, C. Wiencke, A. Wulff, K. Bischof, M. Graeve, C. Wiencke, A. Wulff and K. Bischof (2011) Lipid content and fatty acid consumption in zoospores/developing gametophytes of Saccharina latissima (Laminariales, Phaeophyceae) as potential precursors for secondary metabolites as phlorotannins. Polar Biol. 34, 10111018.
  • 38
    Yates, J. and P. Peckol (1993) Effects of nutrient availability and herbivory on polyphenolic concentrations in the seaweed Fucus vesiculosus. Ecology 74, 17571766.
  • 39
    Abdala-Díaz, R. T., A. Cabello-Pasini, E. Pérez-Rodríguez, R. M. Conde Álvarez and F. L. Figueroa (2005) Daily and seasonal variations of optimum quantum yield and phenolic compounds in Cystoseira tamariscifolia (Phaeophyta). Mar. Biol. 148, 459456.
  • 40
    Pavia, H., G. Cervin, A. Lindgren and P. Åberg (1997) Effects of UV-B radiation and simulated herbivory on phlorotannins in the brown alga Ascophyllum nodosum. Mar. Ecol. Prog. Ser. 157, 139146.
  • 41
    Swanson, A. K. and L. D. Druehl (2002) Induction, exudation and the UV protective role of kelp phlorotannins. Aquat. Bot. 73, 241253.
  • 42
    Schoenwaelder, M., C. Wiencke, M. N. Clayton and K. W. Glombitza (2003) The effect of elevated UV radiation on Fucus spp. (Fucales, Phaeophyta) zygote and embryo development. Plant Biol. 5, 366377.
  • 43
    Connan, S., F. Delisle, E. Deslandes and E. A. Gall (2006) Intra-thallus phlorotannin content and antioxidant activity in Phaeophyceae of temperate waters. Bot. Mar. 49, 3946.
  • 44
    Connan, S., E. Deslandes and E. A. Gall (2007) Influence of day–night and tidal cycles on phenol content and antioxidant capacity in three temperate intertidal brown seaweeds. J. Exp. Mar. Biol. Ecol. 349, 359369.
  • 45
    Ahn, G. N., K. N. Kim, S. H. Cha, C. B. Song, J. Lee, M. S. Heo, I. K. Yeo, N. H. Lee, Y. H. Jee, J. S. Kim, M. S. Heu and Y. J. Jeon (2007) Antioxidant activities of phlorotannins purified from Ecklonia cava on free radical scavenging using ESR and H2O2-mediated DNA damage. Eur. Food Res. Technol. 226, 7179.
  • 46
    Zubia, M., D. Robledo and Y. Freile-Pelegrin (2007) Antioxidant activities in tropical marine macroalgae from the Yucatan Peninsla, Mexico. J. Appl. Phycol. 19, 449458.
  • 47
    Jennings, J. G. and P. D. Steinberg (1994) In situ exudation of phlorotannins by the sublittoral kelp Ecklonia radiate. Mar. Biol. 121, 349354.
  • 48
    Christensen, J. H., B. Hewitson, A. Busuioc, A. Chen, X. Gao, I. Held, R. Jones, R. K. Kolli, W.-T. Kwon, R. Laprise, V. Magaña Rueda, L. Mearns, C. G. Menéndez, J. Räisänen, A. Rinke, A. Sarr and P. Whetton (2007) Regional climate projections. In Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change (Edited by S. Solomon, D. Qin, M. Manning, Z. Chen, M. Marquis, K. B. Averyt, M. Tignor and H. L. Miller), pp. 847940. Cambridge University Press, Cambridge, New York.
  • 49
    McKenzie, R., D. Smale and M. Kotkamp (2004) Relationship between UVB and erythemally weighted radiation. Photochem. Photobiol. Sci. 3, 252256.
  • 50
    Provasoli, L., J. J. A. McLaughlin and M. R. Droop (1957) The development of artificial media for marine algae. Arch. Mikrobiol. 2, 392428.
  • 51
    Koivikko, R., J. Loponen, T. Honkanen and V. Jormalainen (2005) Contents of soluble, cell-wall-bound and exuded phlorotannins in the brown alga Fucus vesiculosus, with implications on their ecological aspects. J. Chem. Ecol. 31, 195212.
  • 52
    Folch, J., M. Lees and G. H. S. Stanley (1957) A simple method for the purification of total lipids from animal tissues. J. Biol. Chem. 226, 497509.
  • 53
    Kattner, G. and H. S. G. Fricke (1986) Simple gas-liquid chromatographic method for the simultaneous determination of fatty acids and alcohols in wax esters of marine organisms. J. Chromatogr. 361, 263268.
  • 54
    R Development Core Team (2011) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. Available at: Accessed on 17 August 2011.
  • 55
    Carroll, R. J. and D. Rupert (1988) Transformation and Weighting in Regression. Chapman and Hall, New York.
  • 56
    Henry, B. E. and K. L. van Alstyne (2004) Effects of UV radiation on growth and phlorotannins in Fucus gardneri (Phaeophyceae) juveniles and embryos. J. Phycol. 40, 527533.
  • 57
    Plouguerné, E., K. Le Lann, S. Connan, G. Jechoux, E. Deslandes and V. Stiger-Pouvreau (2006) Spatial and seasonal variation in density, reproductive status, length and phenolic content of the invasive brown macroalga Sargassum muticum (Yendo) Fensholt along the coast of Western Brittany (France). Aquat. Bot. 85, 377344.
  • 58
    Van Alstyne, K. L., S. L. Whitman and J. M. Ehlig (2001) Differences in herbivore preferences, phlorotannin production, and nutritional quality between juvenile and adult tissues from marine brown algae. Mar. Biol. 139, 201210.
  • 59
    Stiger, V., E. Deslandes and C. E. Payri (2004) Phenolic contents of two brown algae, Turbinaria ornata and Sargassum mangarevense on Tahiti (French Polynesia): Interspecific, ontogenetic and spatio-temporal variations. Bot. Mar. 47, 402409.
  • 60
    Wainman, B. C. and D. R. S. Lean (1992) Carbon fixation into lipid in small freshwater lakes. Limnol. Oceanogr. 37, 956965.
  • 61
    Wainman, B. C., F. R. Pick, P. B. Hamilton and D. R. S. Lean (1993) Lipid production in relation to phytoplankton succession in lakes. Arch. Hydrobiol. 128, 197207.
  • 62
    Khotimchenko, S. V. and I. M. Yakovleva (2004) Effect of solar irradiance on lipids of the green alga Ulva fenestrata Postels et Ruprecht. Bot. Mar. 47, 395401.
  • 63
    Malanga, G. and S. Puntarulo (1995) Oxidative stress and antioxidant content in Chlorella vulgaris after exposure to ultraviolet-B radiation. Physiol. Plant. 94, 672679.
  • 64
    Döhler, G. and I. Biermann (1994) Impact of UV-B radiation on the lipid and fatty acid composition of synchronized Ditylum brightwelli (West) Grunow. Z. Naturforsch. C. 49, 607614.
  • 65
    Thompson, P. A., P. J. Harrison and J. N. C. Whyte (1990) Influence of irradiance on the fatty acid composition of phytoplankton. J. Phycol. 26, 278288.
  • 66
    Poli, G., M. U. Dianzani, K. H. Cheeseman, T. F. Slater, J. Lang and H. Esterbauer (1985) Separation and characterization of the aldehydic products of lipid peroxidation stimulated by carbon tetrachloride or ADP-iron in isolated rat hepatocytes and rat liver microsomal suspensions. Biochem. J. 227, 629638.
  • 67
    DeLange, H. J. and E. van Donk (1997) Effects of UV-B irradiated algae on life history traits of Daphnia pulex. Freshwater Biol. 38, 711720.
  • 68
    Pohl, P. and H. Wagner (1972) Control of fatty acid and lipid biosynthesis in Euglena gracilis by ammonia, light and DCMU. Z. Naturforsch. C 27, 5361.
  • 69
    Lombardy, A. T. and P. J. Wangersky (1991) Influence of phosphorus and silicon on lipid class production by the marine diatom Chaetoceros gracilis grown in turbidostat cage cultures. Mar. Ecol. Prog. Ser. 77, 3947.
  • 70
    Reed, D. C., M. A. Brzezinski, D. A. Coury, W. M. Graham and R. L. Petty (1999) Neutral lipids in macroalgal spores and their role in swimming. Mar. Biol. 133, 737744.
  • 71
    Brzezinski, M. A., D. C. Reed and C. D. Amsler (1993) Neutral lipids as major storage products in zoospores of the giant kelp Macrocystis pyrifera (Phaeophyceae). J. Phycol. 29, 1623.
  • 72
    Floreto, E. A. T., H. Hirata, S. Ando and S. Yamasaki (1993) Effects of temperature, light intensity, salinity and source of nitrogen on the growth, total lipid and fatty acid composition of Ulva pertusa Kjellman (Chlorophyta). Bot. Mar. 36, 149158.
  • 73
    Jamieson, G. R. and E. H. Reid (1972) The component fatty acids of some marine algal lipids. Phytochemistry 11, 14231432.
  • 74
    Mai, K., J. P. Mercer and J. Donlon (1996) Comparative studies on the nutrition of two species of abalone, Haliotis tuberculata L. and Haliotis discus hannai Ino. V. The role of polyunsaturated fatty acids of macroalgae in abalone nutrition. Aquaculture 139, 7789.