Effects of changes in ultraviolet radiation in the Arctic

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Published: February 9, 2010, 3:40 am
Updated: May 7, 2012, 1:03 pm
Author: International Arctic Science Committee
Topic Editor: Mark McGinley
Topics:
This is Section 9.4 of the Arctic Climate Impact Assessment
Lead Author: Harald Loeng; Contributing Authors: Keith Brander, Eddy Carmack, Stanislav Denisenko, Ken Drinkwater, Bogi Hansen, Kit Kovacs, Pat Livingston, Fiona McLaughlin, Egil Sakshaug; Consulting Authors: Richard Bellerby, Howard Browman,Tore Furevik, Jacqueline M. Grebmeier, Eystein Jansen, Steingrimur Jónsson, Lis Lindal Jørgensen, Svend-Aage Malmberg, Svein Østerhus, Geir Ottersen, Koji Shimada

This section assesses the potential impacts of ozone depletion-related increases in solar ultraviolet-B radiation (280–315 nm = UV-B) on arctic marine ecosystems (Effects of changes in ultraviolet radiation in the Arctic). For a comprehensive review of the extensive and rapidly growing technical literature on this subject, readers are referred to several recent books[1], and particularly to Hessen[2] with its focus on the Arctic. UV-B optics in marine waters and ozone layer depletion (Stratospheric Ozone Depletion by Chlorofluorocarbons (Nobel Lecture)) and solar ultraviolet radiation are described in Chapter 5 (Effects of changes in ultraviolet radiation in the Arctic). The exponential relationship between the capacity of ozone to filter ultraviolet light – lower wavelengths are much more strongly filtered – means that small reductions in stratospheric ozone levels result in large increases in UV-B radiation at the earth’s surface[3]. Since ozone layer depletion (Stratospheric Ozone Depletion by Chlorofluorocarbons (Nobel Lecture)) is expected to continue for many more years, albeit at a slower rate[4], the possible impacts of solar UV-B radiation on marine organisms and ecosystems are currently being investigated[5]. A growing number of studies have found that current levels of UVB radiation are harmful to aquatic organisms and may, in some extreme instances, reduce the productivity of marine ecosystems[6]. Reductions in productivity induced by UV-B radiation have been reported for phytoplankton, heterotrophic organisms, and zooplankton; the key intermediary levels of marine food chains[7]. Similar studies on planktonic fish eggs and larvae indicated that exposure to levels of UV-B radiation currently incident at the earth’s surface results in higher mortality and may lead to reduced recruitment success[8].

Ultraviolet radiation also appears to affect biogeochemical cycling within the marine environment and in a manner that could affect overall ecosystem productivity and dynamics[9].

Direct effects on marine organisms (9.4.1)

The majority of UV-B radiation research examines direct effects on specific organisms. Some marine copepods are negatively affected by current levels of UV-B radiation[10]. UV-B-induced mortality in the early life stages, reduced survival and fecundity in females, and changes in sex ratios have all been reported[11]. UV-B-induced damage to the DNA of crustacean zooplankton has also been detected in samples collected up to 20 meters (m) deep[12]. Eggs of Calanus finmarchicus – a prominent member of the mesozooplankton community throughout the North Atlantic – incubated under UV-B radiation exhibited a lower percentage hatch rate than those protected from UV-B radiation[13]. This indicates that Calanus finmarchicus may be sensitive to variation in incident UV-B radiation. Results for the few other species (On the Origin of Species (historical e-book)) that have been studied are highly variable with some showing strong negative impacts, while others are resistant[14]. The factors determining this susceptibility are many and complex, but include seasonality and location of spawning, vertical distribution, presence of UV-B-screening compounds, and the ability to repair UV-B-induced damage to tissues and DNA[15].

The work of Marinaro and Bernard[16], Pommeranz[17], and Hunter et al.[18] provided clear evidence of the detrimental effect of UV-B radiation on the planktonic early life stages of marine fish. Hunter et al.[19], working with northern anchovy (Engraulis mordax) and Pacific mackerel (Scomber japonicus) embryos and larvae, reported that exposure to surface levels of UV-B radiation could be lethal. Significant sub-lethal effects were also reported: lesions in the brain and retina, and reduced growth rate. The study concluded that, under some conditions, 13% of the annual production of northern anchovy larvae could be lost as a result of UV-B-related mortality[20]. Atlantic cod eggs were negatively affected by exposure to UV-B radiation in very shallow water; 50 cm deep or less[21]. With the exception of a small (but rapidly growing) number of recent studies, little additional information is available on the effects of UV-B radiation on the early life stages of fish. However, as for copepods, the early life stages of fish will vary in their susceptibility to UV-B radiation and for the same reasons. Thus, some studies conclude that the effects of UV-B radiation will be significant[22], while others conclude that they will not[23].

Indirect effects on marine organisms (9.4.2)

Exposure to UV radiation, especially UV-B radiation, has many harmful effects on health. These may result in poorer performance, or even death, despite not being directly induced by exposure to UV-B radiation. UV-B radiation suppresses systemic and local immune responses to a variety of antigens, including microorganisms[24]. In addition to suppressing T-cell-mediated immune reactions, UV-B radiation also affects nonspecific cellular immune defenses. Recent studies demonstrate disturbed immunological responses in UV-B-irradiated roach (Rutilus rutilus): the function of isolated head kidney neutrophils and macrophages (immuno-responsive cells) were significantly altered after a single dose of UV-B radiation[25]. Natural cytotoxicity, assumed to be an important defense mechanism in viral, neoplastic, and parasitic diseases, was also reduced. A single dose of UV-B radiation exposure decreased the ability of fish lymphocytes to respond to activators, and this was still apparent 14 days later[26]. This indicates altered regulation of lymphocyte-dependent immune functions. Finally, exposure to UV-B radiation induces a strong systemic stress response which is manifested in fish blood by an increased number of circulating phagocytes and elevated plasma cortisol levels[27]. Exposure to UV-A (315–400 nm) radiation induced some of the same negative effects on the immune system[28]. Since high cortisol levels induce immunosuppression in fish[29] the effect of exposure to UV-B radiation on the immune system clearly has both direct and indirect components. Taken together, these findings indicate that the immune system of fish is significantly affected by exposure to a single, moderate-level dose of UV-B radiation. At the population level, a reduction in immune response might be manifested as lowered resistance to pathogens and increased susceptibility to disease. The ability of the fish immune system to accommodate increases in solar UV-B radiation is not known. Also, the immune system of young fish is likely to be highly vulnerable to UV-B radiation because lymphoid organs are rapidly developing and because critical phases of cell proliferation, differentiation, and maturation are occurring[30]. It is also possible that exposure to ambient UV-B radiation impedes the development of the thymus or other lymphoid organs resulting in compromised immune defense later in life. The effect of UV-B radiation on the immune function of fish embryos and larvae, and on the development of the immune system, is unknown.

Other indirect effects of UV-B radiation are also possible. For example, UV-B radiation may affect sperm quality for species that spawn in the surface layer[31] and so affect fertilization rate and/or genome transfer.

Studies on the impact of UV-B radiation have almost all examined the effects of short-term exposure on biological end-points such as skin injury (sunburn), DNA damage, development and growth rates, immune function, or outright mortality. Few have examined the potential effects of longer-term (low-level) exposure[32]. All these indirect (and/or longer-term) effects of UV-B radiation have yet to be investigated.

Ecosystem effects (9.4.3)

Food chains (9.4.3.1)

Fig. 9.32. Output of a mathematical simulation model[33] illustrating the relative effects of selected variables on UV-induced mortality in Calanus finmarchicus embryos[34]. The plots illustrate the effects on irradiance of (a) clear versus cloudy sky, (b) clear versus an opaque water column, (c) 50% thinning of ozone versus ambient ozone, while (d) compares the relative impacts of all three on mortality. This graphic illustrates that water column transparency is the single most important determinant of UV exposure – of considerably more importance than ozone layer depletion.

Although the effects of UV-B radiation are strongly species-specific, marine bacterioplankton and phytoplankton can be negatively affected[35]. Severe exposure to UV-B radiation can, therefore, decrease productivity at the base of marine food chains. The importance of this decrease is highly speculative, but decreases in carbon fixation of 20 to 30% have been proposed[36]. Arctic phytoplankton appear more susceptible than antarctic species, possibly owing to deeper surface mixed layers in the Arctic[37]. Also, if UV-B radiation reduces the productivity of protozoans and crustacean zooplankton there will be less prey available for fish larvae and other organisms that feed upon them. The few studies that have investigated the indirect effects of UV-B radiation on specific organisms conclude that UV-B-induced changes in food-chain interactions can be far more significant than direct effects on individual organisms at any single trophic level[38]. Recent investigations indicate the possibility of food-chain effects in both the marine and freshwater environment: exposure to UV-B radiation (even at low dose rates) reduces the total lipid content of some microalgae[39] and this includes the polyunsaturated fatty acids (PUFAs)[40]. For zooplankton and fish larvae, the only source of PUFAs is the diet – they cannot be synthesized and so must be obtained from prey organisms[41]. Dietary deficiencies are manifested in many ways. For example, in the freshwater cladoceran Daphnia spp., growth rates are correlated with the concentration of eicosapentaenoic acid in the water column[42]. In Atlantic herring, dietary deficiencies of essential fatty acids, in particular docosahexaenoic acid, reduce the number of rods in the eyes[43] and negatively affect feeding at low light levels[44]. Other negative consequences of essential fatty acid deficits have also been reported[45]. A UV-B-induced reduction in the PUFA content of microalgae will be transferred to the herbivorous zooplankton that graze on them, thereby decreasing the availability of this essential fatty acid to fish larvae. Since fish larvae (and their prey) require these essential fatty acids for proper development and growth, a reduction in the nutritional quality of the food base has potentially widespread and significant implications for the overall productivity and health of aquatic ecosystems.

Quantitative assessments (9.4.3.2)

Quantitative assessments of the effects of UV-B radiation on marine organisms at the population level are scarce. However, several studies are currently underway using mathematical simulation models. Neale et al.[46] estimated that a 50% seasonal reduction in stratospheric ozone levels could reduce total levels of primary production – integrated throughout the water column – by up to 8.5%. Kuhn et al.[47] developed a model that incorporates physical and biological information and were able to generate an absolute estimate of mortality under different meteorological and hydrographic conditions. As a result, they were able to evaluate the relative impacts of different combinations of environmental conditions – for example, a typical clear sky versus a typical overcast sky; a typical clear water column versus a typical opaque coastal water column; current ambient ozone levels versus a realistically thinned ozone layer. For Calanus finmarchicus eggs in the estuary and Gulf of St. Lawrence, UV-B-induced mortality for all model scenarios ranged from <1% to 51%, with a mean of 10.05% and an uncertainty of ±11.9% (based on 1 standard deviation and 48 modeled scenarios). For Atlantic cod, none of the scenarios gave a UV-B-induced mortality greater than 1.2%, and the mean was 1.0±0.63% (72 modeled scenarios).

In both assessments[48], the most important determinant of UV-B related effects was water column transparency (see Fig. 9.32): even when ozone layer depletions of 50% were modeled, the effect on mortality remained far lower than that resulting from either thick cloud cover or opacity of the water column. This demonstrates that variability in cloud cover, water quality, and vertical distribution and displacement within the surface mixed layer have a greater effect on the flux of UV-B radiation to which planktonic marine organisms are exposed than ozone layer depletion. In contrast, Huot et al.[49] showed that ozone thickness could in some instances be the single most important determinant of DNA damage in bacterioplankton.

Fig. 9.33. Level of protection from UV damage afforded by the organic matter content of the water column. (a) diffuse attenuation coefficient (Kd) at 305 nm versus modeled survival of Atlantic cod embryos exposed to UV radiation in a mixed water column; (b) Kd at 305 nm versus modeled survival of Calanus finmarchicus embryos exposed to UV radiation in a mixed water column; (c) dissolved organic carbon (DOC) versus Kd at 305 nm from field measurements in temperate marine coastal waters (the estuary and Gulf of St. Lawrence, Canada). The straight line is the regression; the curved lines the 95% confidence intervals[50].

Since the concentrations of dissolved organic carbon (DOC) and Chl-a are strongly correlated with the transparency of the water column to UV-B radiation, it follows that their concentrations are an overriding factor affecting UV-B-induced mortality. The Kuhn et al.[51] model supports this contention. DOC levels in eutrophic coastal zones are often greater than 3 to 4 mg/L; the diffuse attenuation coefficients for UV-B radiation at such levels essentially protect Calanus finmarchicus and cod eggs from UV-B-induced mortality (Fig. 9.33). Thus, DOC can be considered as a sunscreen for organisms inhabiting eutrophic coastal zone waters. DOC concentrations in arctic waters are typically <1 mg/L[52]. At these levels, DOC is not as effective at protecting planktonic marine organisms from UV-B-related damage (Fig. 9.33).

Although these model-based predictions are useful, there are limited data to parameterize the models, and it will be some time before similar predictions can be made for the many species inhabiting the full range of conditions within the world’s ocean, including those of the Arctic.

General perspectives (9.4.4)

Although UV-B radiation can have negative impacts (direct effects) on marine organisms and [[population]s], it is only one of many environmental factors (e.g., bacterial and/or viral pathogens, predation, toxic algae) that result in the mortality typically observed in these organisms. Recent assessments indicate that UV-B radiation is generally only a minor source of direct mortality (or decreases in productivity) for populations, particularly in "DOC-protected" coastal zones. However, for those species whose early life stages occur near the surface, there may be circumstances (albeit rare) –such as a cloudless sky, thin ozone layer, lack of wind, calm seas, low nutrient loading – under which the contribution of UV-B radiation to the productivity and/or mortality of a population could be far more significant. The impact of indirect effects has not as yet been adequately evaluated.

Chapter 9: Marine Systems
9.1. Introduction (Effects of changes in ultraviolet radiation in the Arctic)
9.2. Physical oceanography
9.2.1. General features (Effects of changes in ultraviolet radiation in the Arctic)
9.2.2. Sea ice (Sea ice effect on marine systems in the Arctic)
9.2.3. Ocean processes of climatic importance
9.2.4. Variability in hydrographic properties and currents
9.2.5. Anticipated changes in physical conditions
9.3. Biota
9.3.1. General description of the Arctic biota community
9.3.2. Physical factors mediating ecological change
9.3.3. Past variability – interannual to decadal
9.3.4. Future change – processes and impacts on biota
9.4. Effects of changes in ultraviolet radiation
9.5. The carbon cycle and climate change
9.6. Key findings (Effects of changes in ultraviolet radiation in the Arctic)
9.7. Gaps in knowledge and research needs

References

Citation

Committee, I. (2012). Effects of changes in ultraviolet radiation in the Arctic. Retrieved from http://editors.eol.org/eoearth/wiki/Effects_of_changes_in_ultraviolet_radiation_in_the_Arctic
  1. ↑ De Mora, S.J., S. Demers and M. Vernet (eds.), 2000. The Effects of UV Radiation in the Marine Environment. Cambridge University Press.–Häder, D.-P. (ed.), 1997. The Effects of Ozone Depletion on Aquatic Ecosystems. R.G. Landes Company, Austin, Texas.–Helbling, E.W. and H. Zagarese, 2003. UV Effects in Aquatic Organisms and Ecosystems. Springer-Verlag. 574pp.
  2. ↑ Hessen, D.O. (ed.), 2001. UV Radiation and Arctic Ecosystems. Springer-Verlag. 310pp.
  3. ↑ Kerr, J.B. and C.T. McElroy, 1993. Evidence for large upward trends of ultraviolet-B radiation linked to ozone depletion. Science, 262:1032–1034.–Madronich, S., R.L. McKenzie, M.M. Caldwell and L.O. Bjorn, 1995. Changes in ultraviolet radiation reaching the Earth's surface. Ambio, 24:143–152.
  4. ↑ Shindell, D.T., D. Rind and P. Lonergan, 1998. Increased polar stratospheric ozone losses and delayed eventual recovery owing to increasing greenhouse-gas concentrations. Nature, 392:589–592.–Staehelin, J., N.R.P. Harris, C. Appenzeller and J. Eberhard, 2001. Ozone trends: a review. Reviews of Geophysics, 39:231–290.–Taalas, P., J. Kaurola, A. Kylling, D. Shindell, R. Sausen, M. Dameris, V. Grewe, J. Herman, J. Damski and B. Steil, 2000. The impact of greenhouse gases and halogenated species on future solar UV radiation doses. Geophysical Research Letters, 27:1127–1130.
  5. ↑ Browman, H.I., 2003. Assessing the impacts of solar ultraviolet radiation on the early life stages of crustacean zooplankton and ichthyoplankton in marine coastal systems. Estuaries, 26:30–39.–Browman, H.I., C. Alonso Rodriguez, F. Beland, J.J. Cullen, R.F. Davis, J.H.M. Kouwenberg, P.S. Kuhn, B. McArthur, J.A. Runge, J.-F. St-Pierre and R.D. Vetter, 2000. Impact of ultraviolet radiation on marine crustacean zooplankton and ichthyoplankton: a synthesis of results from the estuary and Gulf of St. Lawrence, Canada. Marine Ecology Progress Series, 199:293–311.–De Mora, S.J., S. Demers and M. Vernet (eds.), 2000. The Effects of UV Radiation in the Marine Environment. Cambridge University Press.–Häder, D.-P. (ed.), 1997. The Effects of Ozone Depletion on Aquatic Ecosystems. R.G. Landes Company, Austin, Texas.–Häder, D.-P., H.D. Kumar, R.C. Smith and R.C. Worrest, 2003. Aquatic ecosystems: effects of solar ultraviolet radiation and interactions with other climatic change factors. Photochemical and Photobiological Sciences, 2:39–50.–Helbling, E.W. and H. Zagarese, 2003. UV Effects in Aquatic Organisms and Ecosystems. Springer-Verlag. 574pp.–Hessen, D.O. (ed.), 2001. UV Radiation and Arctic Ecosystems. Springer-Verlag. 310pp.
  6. ↑ De Mora, S.J., S. Demers and M. Vernet (eds.), 2000. The Effects of UV Radiation in the Marine Environment. Cambridge University Press.–Häder, D.-P. (ed.), 1997. The Effects of Ozone Depletion on Aquatic Ecosystems. R.G. Landes Company, Austin, Texas.–Häder, D.-P., H.D. Kumar, R.C. Smith and R.C. Worrest, 2003. Aquatic ecosystems: effects of solar ultraviolet radiation and interactions with other climatic change factors. Photochemical and Photobiological Sciences, 2:39–50.–Helbling, E.W. and H. Zagarese, 2003. UV Effects in Aquatic Organisms and Ecosystems. Springer-Verlag. 574pp. Hessen, D.O. (ed.), 2001. UV Radiation and Arctic Ecosystems. Springer-Verlag. 310pp.–Hessen, D.O. (ed.), 2001. UV Radiation and Arctic Ecosystems. Springer-Verlag. 310pp.
  7. ↑ De Mora, S.J., S. Demers and M. Vernet (eds.), 2000. The Effects of UV Radiation in the Marine Environment. Cambridge University Press.–Häder, D.-P. (ed.), 1997. The Effects of Ozone Depletion on Aquatic Ecosystems. R.G. Landes Company, Austin, Texas.–Häder, D.-P., H.D. Kumar, R.C. Smith and R.C. Worrest, 2003. Aquatic ecosystems: effects of solar ultraviolet radiation and interactions with other climatic change factors. Photochemical and Photobiological Sciences, 2:39–50.–Helbling, E.W. and H. Zagarese, 2003. UV Effects in Aquatic Organisms and Ecosystems. Springer-Verlag. 574pp. Hessen, D.O. (ed.), 2001. UV Radiation and Arctic Ecosystems. Springer-Verlag. 310pp.–Hessen, D.O. (ed.), 2001. UV Radiation and Arctic Ecosystems. Springer-Verlag. 310pp.
  8. ↑ Hunter, J.R., S.E. Kaupp and J.H. Taylor, 1981. Effects of solar and artificial ultraviolet-B radiation on larval northern anchovy, Engraulis mordax. Photochemistry and Photobiology, 34:477–486.–Hunter, J.R., S.E. Kaupp and J.H. Taylor, 1982. Assessment of effects of UV radiation on marine fish larvae. In: J. Calkins (ed.). The Role of Solar Ultraviolet Radiation in Marine Ecosystems, pp. 459–493. Plenum Press.–Lesser, M.P., J.H. Farrell and C.W. Walker, 2001. Oxidative stress, DNA damage and p53 expression in the larvae of Atlantic cod (Gadus morhua) exposed to ultraviolet (290–400 nm) radiation. Journal of Experimental Biology, 204:157–164.–Pommeranz, T., 1974. Resistance of plaice eggs to mechanical stress and light. In: J.H.S. Blaxter (ed.). The Early Life History of Fish, pp. 397–416. Springer-Verlag.–Walters, C. and B. Ward, 1998. Is solar radiation responsible for declines in marine survival rates of anadromous salmonids that rear in small streams? Canadian Journal of Fisheries and Aquatic Sciences, 55:2533–2538.–Williamson, C.E., S.L. Metzgar, P.A. Lovera and R.E. Moeller, 1997. Solar ultraviolet radiation and the spawning habitat of yellow perch, Perca flavescens. Ecological Applications, 7:1017–1023.–Zagarese, H.E. and C.E. Williamson, 2000. Impact of solar UV radiation on zooplankton and fish. In: S.J. de Mora, S. Demers and M. Vernet (eds.). The Effects of UV Radiation in the Marine Environment, pp. 279–309. Cambridge University Press.–Zagarese, H.E. and C.E. Williamson, 2001. The implications of solar UV radiation exposure for fish and fisheries. Fish and Fisheries, 2:250–260.
  9. ↑ Zepp, R.G., T.V. Callaghan and D.J. Erickson III, 2003. Interactive effects of ozone depletion and climate change on biogeochemical cycles. Photochemical and Photobiological Sciences, 2:51–61.
  10. ↑ Häder, D.-P., H.D. Kumar, R.C. Smith and R.C. Worrest, 2003. Aquatic ecosystems: effects of solar ultraviolet radiation and interactions with other climatic change factors. Photochemical and Photobiological Sciences, 2:39–50.
  11. ↑ Chalker-Scott, L., 1995. Survival and sex ratios of the intertidal copepod, Tigriopus californicus, following ultraviolet-B (290–320 nm) radiation exposure. Marine Biology, 123:799–804.–Karanas, J.J., H. Van Dyke and R.C. Worrest, 1979. Midultraviolet (UV-B) sensitivity of Acartia clausii Giesbrecht (Copepoda). Limnology and Oceanography, 24:1104–1116.–Karanas, J.J., R.C. Worrest and H. Van Dyke, 1981. Impact of UV-B radiation on the fecundity of the copepod Acartia clausii. Marine Biology, 65:125–133.–Lacuna, D.G. and S.-I. Uye, 2001. Influence of mid-ultraviolet (UVB) radiation on the physiology of the marine planktonic copepod Acartia omorii and the potential role of photoreactivation. Journal of Plankton Research, 23:143–156.–Naganuma, T., T. Inoue and S. Uye, 1997. Photoreactivation of UV-induced damage to embryos of a planktonic copepod. Journal of Plankton Research, 19:783–787.–Tartarotti, B., W. Cravero and H.E. Zagarese, 2000. Biological weighting function for the mortality of Boeckella gracilipes (Copepoda, Crustacea) derived from experiments with natural solar radiation. Photochemistry and Photobiology, 72:314–319.
  12. ↑ Malloy, K.D., M.A. Holman, D. Mitchell and H.W. Detrich III, 1997. Solar UVB-induced DNA damage and photoenzymatic DNA repair in Antarctic zooplankton. Proceedings of the National Academy of Sciences, 94:1258–1263.
  13. ↑ Alonso Rodriguez, C., H.I. Browman, J.A. Runge and J.-F. St-Pierre, 2000. Impact of solar ultraviolet radiation on hatching of a marine copepod, Calanus finmarchicus. Marine Ecology Progress Series, 193:85–93.
  14. ↑ Damkaer, D.M., 1982. Possible influences of solar UV radiation in the evolution of marine zooplankton. In: J. Calkins (ed.). The Role of Solar Ultraviolet Radiation in Marine Ecosystems, pp. 701–706. Plenum Press.–Dey, D.B., D.M. Damkaer and G.A. Heron, 1988. UV-B dose/dose-rate responses of seasonally abundant copepods of Puget Sound. Oecologia, 76:321–329.–Thomson, B.E., 1986. Is the impact of UV-B radiation on marine zooplankton of any significance? In: J.G. Titus (ed.). Effects of Changes in Stratospheric Ozone and Global Climate, Vol. 2, pp. 203–209. US Environmental Protection Agency and United Nations Environment Programme.–Zagarese, H.E. and C.E.Williamson, 2001. The implications of solar UV radiation exposure for fish and fisheries. Fish and Fisheries, 2:250–260.
  15. ↑ Williamson, C.E., P.J. Neale, G. Grad, H.J. De Lange and B.R. Hargreaves, 2001. Beneficial and detrimental effects of UV on aquatic organisms: Implications of spectral variation. Ecological Applications, 11:1843–1857.
  16. ↑ Marinaro, J. and M. Bernard, 1966. Contribution à l'étude des oeufs et larves pélagiques de poissons méditerranés. I. Notes preliminaires sur l'influence léthale du rayonnement solaire sur les oeufs. Pelagos, 6:49–55.
  17. ↑ Pommeranz, T., 1974. Resistance of plaice eggs to mechanical stress and light. In: J.H.S. Blaxter (ed.).The Early Life History of Fish, pp. 397–416. Springer-Verlag.
  18. ↑ Hunter, J.R., J.H. Taylor and H.G. Moser, 1979. Effect of ultraviolet irradiation on eggs and larvae of the northern anchovy, Engraulis mordax, and the Pacific mackerel, Scomber japonicus, during the embryonic stage. Photochemistry and Photobiology, 29:325–338.–Hunter, J.R., S.E. Kaupp and J.H. Taylor, 1981. Effects of solar and artificial ultraviolet-B radiation on larval northern anchovy, Engraulis mordax. Photochemistry and Photobiology, 34:477–486.–Hunter, J.R., S.E. Kaupp and J.H. Taylor, 1982. Assessment of effects of UV radiation on marine fish larvae. In: J. Calkins (ed.). The Role of Solar Ultraviolet Radiation in Marine Ecosystems, pp. 459–493. Plenum Press.
  19. ↑ Hunter, J.R., J.H. Taylor and H.G. Moser, 1979. Effect of ultraviolet irradiation on eggs and larvae of the northern anchovy, Engraulis mordax, and the Pacific mackerel, Scomber japonicus, during the embryonic stage. Photochemistry and Photobiology, 29:325–338.
  20. ↑ Hunter, J.R., S.E. Kaupp and J.H. Taylor, 1981. Effects of solar and artificial ultraviolet-B radiation on larval northern anchovy, Engraulis mordax. Photochemistry and Photobiology, 34:477–486.–Hunter, J.R., S.E. Kaupp and J.H. Taylor, 1982. Assessment of effects of UV radiation on marine fish larvae. In: J. Calkins (ed.). The Role of Solar Ultraviolet Radiation in Marine Ecosystems, pp. 459–493. Plenum Press.
  21. ↑ Béland, F., H.I. Browman, C. Alonso Rodriguez and J.-F. St-Pierre, 1999. Effect of solar ultraviolet radiation (280–400 nm) on the eggs and larvae of Atlantic cod (Gadus morhua). Canadian Journal of Fisheries and Aquatic Sciences, 56:1058–1067.–Browman, H.I. and R.D. Vetter, 2001. Impact of UV radiation on crustacean zooplankton and ichthyoplankton: case studies from sub-arctic marine ecosystems. In: D.O. Hessen (ed.). UV Radiation and Arctic Ecosystems, pp. 261–304. Springer-Verlag.
  22. ↑ Battini, M., V. Rocco, M. Lozada, B. Tartarotti and H.E. Zagarese, 2000. Effects of ultraviolet radiation on the eggs of landlocked Galaxias maculatus (Galaxiidae, Pisces) in northwestern Patagonia. Freshwater Biology, 44:547–552.–Lesser, M.P., J.H. Farrell and C.W. Walker, 2001. Oxidative stress, DNA damage and p53 expression in the larvae of Atlantic cod (Gadus morhua) exposed to ultraviolet (290–400 nm) radiation. Journal of Experimental Biology, 204:157–164.–Williamson, C.E., S.L. Metzgar, P.A. Lovera and R.E. Moeller, 1997. Solar ultraviolet radiation and the spawning habitat of yellow perch, Perca flavescens. Ecological Applications, 7:1017–1023.
  23. ↑ Dethlefsen, V., H. von Westernhagen, H. Tug, P.D. Hansen and H. Dizer, 2001. Influence of solar ultraviolet-B on pelagic fish embryos: osmolality, mortality and viable hatch. Helgoland Marine Research, 55:45–55.–Kuhn, P.S., H.I. Browman, R.F. Davis, J.J. Cullen and B.L. McArthur, 2000. Modeling the effects of ultraviolet radiation on embryos of Calanus finmarchicus and Atlantic cod (Gadus morhua) in a mixing environment. Limnology and Oceanography, 45:1797–1806.–Steeger, H.U., J.F. Freitag, S. Michl, M. Wiemer and R.J. Paul, 2001. Effects of UV-B radiation on embryonic, larval and juvenile stages of North Sea plaice (Pleuronectes platessa) under simulated ozonehole conditions. Helgoland Marine Research, 55:56–66.
  24. ↑ Garssen, J., M. Norval, A. El-Ghorr, N.K. Gibbs, C.D. Jones, D. Cerimele, C. De Simone, S. Caffieri, F. Dall'Acqua, F.R. De Gruijl, Y. Sontag and H. Van Loveren, 1998. Estimation of the effect of increasing UVB exposure on the human immune system and related resistance to infectious diseases and tumours. Journal of Photochemistry and Photobiology B, 42:167–179.–Hurks, M., J. Garssen, H. van Loveren and B.-J. Vermeer, 1994. General aspects of UV-irradiation on the immune system. In: G. Jori, R.H. Pottier, M.A.J. Rodgers and T.G. Truscott (eds.). Photobiology in Medicine, pp. 161–176. Plenum Press.
  25. ↑ Salo, H.M., T.M. Aaltonen, S.E. Markkula and E.I. Jokinen, 1998. Ultraviolet B irradiation modulates the immune system of fish (Rutilus rutilus, Cyprinidae). I. Phagocytes. Journal of Photochemistry and Photobiology, 67:433–437.
  26. ↑ Jokinen, E.I., H.M. Salo, S.E. Markkula, A.K. Immonen and T.M. Aaltonen, 2001. Ultraviolet B irradiation modulates the immune system of fish (Rutilus rutilus, Cyprinidae). Part III. Lymphocytes. Photochemistry and Photobiology, 73:505–512.
  27. ↑ Salo, H.M., E.I. Jokinen, S.E. Markkula and T.M. Aaltonen, 2000a. Ultraviolet B irradiation modulates the immune system of fish (Rutilus rutilus, Cyprinidae). II. Blood. Journal of Photochemistry and Photobiology, 71:65–70.
  28. ↑ Salo, H.M., E.I. Jokinen, S.E. Markkula , T.M. Aaltonen and H.T. Penttila, 2000b. Comparative effects of UVA and UVB irradiation on the immune system of fish. Journal of Photochemistry and Photobiology B, 56:154–162.
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  30. ↑ Botham, J.W. and M.J. Manning, 1981. The histogenesis of the lymphoid organs in the carp Cyprinus carpio L. and the ontogenetic development of allograft reactivity. Journal of Fish Biology, 19:403–414.–Chilmonczyk, S., 1992. The thymus in fish: Development and possible function in the immune response. Annual Review of Fish Diseases, 2:181–200.–Grace, M.F. and M.J. Manning, 1980. Histogenesis of the lymphoid organs in rainbow trout, Salmo gairdneri. Developmental and Comparative Immunology, 4:255–264.
  31. ↑ Don, J. and R.R. Avtalion, 1993. Ultraviolet irradiation of tilapia spermatozoa and the Hertwig effect: electron microscopic analysis. Journal of Fish Biology, 42:1–14.–Valcarcel, A., G. Guerrero and M.C. Maggese, 1994. Hertwig effect caused by UV-irradiation of sperm of the catfish, Rhamdia sapo (Pisces, Pimelodidae), and its photoreactivation. Aquaculture, 128:21–28.
  32. ↑ Fidhiany, L. and K. Winckler, 1999. Long term observation on several growth parameters of convict cichlid under an enhanced ultraviolet-A (320–400 nm) irradiation. Aquaculture International, 7:1–12.
  33. ↑ Kuhn, P.S., H.I. Browman, R.F. Davis, J.J. Cullen and B.L. McArthur, 2000. Modeling the effects of ultraviolet radiation on embryos of Calanus finmarchicus and Atlantic cod (Gadus morhua) in a mixing environment. Limnology and Oceanography, 45:1797–1806.
  34. ↑ Browman, H.I., C. Alonso Rodriguez, F. Beland, J.J. Cullen, R.F. Davis, J.H.M. Kouwenberg, P.S. Kuhn, B. McArthur, J.A. Runge, J.-F. St-Pierre and R.D. Vetter, 2000. Impact of ultraviolet radiation on marine crustacean zooplankton and ichthyoplankton: a synthesis of results from the estuary and Gulf of St. Lawrence, Canada. Marine Ecology Progress Series, 199:293–311.
  35. ↑ De Mora, S.J., S. Demers and M. Vernet (eds.), 2000. The Effects of UV Radiation in the Marine Environment. Cambridge University Press.–Hessen, D.O. (ed.), 2001. UV Radiation and Arctic Ecosystems. Springer-Verlag. 310pp.
  36. ↑ Helbling, E.W. and V.E. Villafañe, 2001. UV radiation effects on phytoplankton primary production: a comparison between Arctic and Antarctic marine ecosystems. In: D.O. Hessen (ed.). UV Radiation and Arctic Ecosystems, pp. 203–226. Springer-Verlag.
  37. ↑ Helbling, E.W. and V.E. Villafañe, 2001. UV radiation effects on phytoplankton primary production: a comparison between Arctic and Antarctic marine ecosystems. In: D.O. Hessen (ed.). UV Radiation and Arctic Ecosystems, pp. 203–226. Springer-Verlag.
  38. ↑ Bothwell, M.L., D.M.J. Sherbot and C.M. Pollock, 1994. Ecosystem response to solar ultraviolet-B radiation: influence of trophic-level interactions. Science, 265:97–100.–Hessen, D.O., H.J. De Lange and E. Van Donk, 1997. UV-induced changes in phytoplankton cells and its effects on grazers. Freshwater Biology, 38:513–524.–Williamson, C.E., B.R. Hargreaves, P.S. Orr and P.A. Lovera, 1999. Does UV play a role in changes in predation and zooplankton community structure in acidified lakes– Limnology and Oceanography, 44:774–783.
  39. ↑ Arts, M.T. and H. Rai, 1997. Effects of enhanced ultraviolet-B radiation on the production of lipid, polysaccharide and protein in three freshwater algal species. Freshwater Biology, 38:597–610.–Arts, M.T., H. Rai and V.P. Tumber, 2000. Effects of artificial UV-A and UV-B radiation on carbon allocation in Synechococcus elongatus (cyanobacterium) and Nitzschia palea (diatom).Verhandlungen der Internationalen Vereinigung fur Theoretische und Angewandte Limnologie, 27:2000–2007.–Plante, A.J. and M.T. Arts, 1998. Photosynthate production in laboratory cultures (UV conditioned and unconditioned) of Cryptomonas erosa under simulated doses of UV radiation. Aquatic Ecology, 32:297–312.
  40. ↑ Goes, J.I., N. Handa, S. Taguchi and T. Hama, 1994. Effect of UV-B radiation on the fatty acid composition of the marine phytoplankton Tetraselmis sp.: relationship to cellular pigments. Marine Ecology Progress Series, 114:259–274.–Hessen, D.O., H.J. De Lange and E. Van Donk, 1997. UV-induced changes in phytoplankton cells and its effects on grazers. Freshwater Biology, 38:513–524.–Wang, K.S. and T. Chai, 1994. Reduction in omega-3 fatty acids by UV-B irradiation in microalgae. Journal of Applied Phycology, 6:415–421.
  41. ↑ Goulden, C.E. and A.R. Place, 1990. Fatty acid synthesis and accumulation rates in daphnids. Journal of Experimental Zoology, 256:168–178.–Rainuzzo, J.R., K.I. Reitan and Y. Olsen, 1997. The significance of lipids at early stages of marine fish: a review. Aquaculture, 155:103–115.–Reitan, K.I., J.R. Rainuzzo, G. Øie and Y. Olsen, 1997. A review of the nutritional effects of algae in marine fish larvae. Aquaculture, 155:207–221.–Sargent, J.R., L.A. McEvoy and J.G. Bell, 1997. Requirements, presentation and sources of polyunsaturated fatty acids in marine fish larval feeds. Aquaculture, 155:117–127.
  42. ↑ De Lange, H.J. and E. Van Donk, 1997. Effects of UVB-irradiated algae on life history traits of Daphnia pulex. Freshwater Biology, 38:711–720.–Müller-Navarra, D., 1995a. Evidence that a highly unsaturated fatty acid limits Daphnia growth in nature. Archives of Hydrobiology, 132:297–307.–Müller-Navarra, D., 1995b. Biochemical versus mineral limitation in Daphnia. Limnology and Oceanography, 40:1209–1214.–Scott, C.L., S. Falk-Petersen, J.R. Sargent, H. Hop, O.J. Lønne and M. Poltermann, 1999. Lipids and trophic interactions of ice fauna and pelagic zooplankton in the marginal ice zone of the Barents Sea. Polar Biology, 21:65–70.
  43. ↑ Bell, M.V. and J.R. Dick, 1993. The appearance of rods in the eyes of herring and increased didocosahexaenoyl molecular species of phospholipids. Journal of the Marine Biological Association of the UK, 73:679–688.
  44. ↑ Bell, M.V., R.S. Batty, J.R. Dick, K. Fretwell, J.C. Navarro and J.R. Sargent, 1995. Dietary deficiency of docosahexaenoic acid impairs vision at low light intensities in juvenile herring (Clupea harengus L.). Lipids, 30:443–449.–Masuda, R., T. Takeuchi, K. Tsukamoto, Y. Ishizaki, M. Kanematsu and K. Imaizumi, 1998. Critical involvement of dietary docosahexaenoic acid in the ontogeny of schooling behaviour in the yellowtail. Journal of Fish Biology, 53:471–484.
  45. ↑ Bell, J.G., D.R. Tocher, B.M. Farndale and J.R. Sargent, 1998. Growth, mortality, tissue histopathology and fatty acid compositions, eicosanoid production and response to stress, in juvenile turbot fed diets rich in gamma-linolenic acid in combination with eicosapentaenoic acid or docosahexaenoic acid. Prostaglandins, Leukotrienes and Essential Fatty Acids, 58:353–364.–Kanazawa, A., 1997. Effects of docosahexaenoic acid and phospholipids on stress tolerance of fish. Aquaculture, 155:129–134.–Rainuzzo, J.R., K.I. Reitan and Y. Olsen, 1997.The significance of lipids at early stages of marine fish: a review. Aquaculture, 155:103–115.
  46. ↑ Neale, P.J., R.F. Davis and J.J. Cullen, 1998. Interactive effects of ozone depletion and vertical mixing on photosynthesis of Antarctic phytoplankton. Nature, 392:585–589.–Neale, P.J., J.J. Fritz and R.F. Davis, 2001. Effects of UV on photosynthesis of Antarctic phytoplankton: models and their application to coastal and pelagic assemblages. Revista Chilena de Historia Natural, 74:283–292.
  47. ↑ Kuhn, P.S., H.I. Browman, R.F. Davis, J.J. Cullen and B.L. McArthur, 2000. Modeling the effects of ultraviolet radiation on embryos of Calanus finmarchicus and Atlantic cod (Gadus morhua) in a mixing environment. Limnology and Oceanography, 45:1797–1806.
  48. ↑ Kuhn, P.S., H.I. Browman, R.F. Davis, J.J. Cullen and B.L. McArthur, 2000. Modeling the effects of ultraviolet radiation on embryos of Calanus finmarchicus and Atlantic cod (Gadus morhua) in a mixing environment. Limnology and Oceanography, 45:1797–1806.Kuhn et al., 2000; Neale et al., 1998, 2001
  49. ↑ Huot, Y., W.H. Jeffrey, R.F. Davis and J.J. Cullen, 2000. Damage to DNA in bacterioplankton:A model of damage by ultraviolet radiation and its repair as influenced by vertical mixing. Photochemistry and Photobiology, 72:62–74.
  50. ↑ Browman, H.I., 2003. Assessing the impacts of solar ultraviolet radiation on the early life stages of crustacean zooplankton and ichthyoplankton in marine coastal systems. Estuaries, 26:30–39.
  51. ↑ Kuhn, P.S., H.I. Browman, R.F. Davis, J.J. Cullen and B.L. McArthur, 2000. Modeling the effects of ultraviolet radiation on embryos of Calanus finmarchicus and Atlantic cod (Gadus morhua) in a mixing environment. Limnology and Oceanography, 45:1797–1806.
  52. ↑ Aas, E., J. Høkedal, N.K. Højerslev, R. Sandvik and E. Sakshaug, 2001. Spectral properties and UV attenuation in Arctic marine waters. In: D.O. Hessen (ed.). UV Radiation and Arctic Ecosystems, pp. 23–56. Springer-Verlag.
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