Effects of changes in ultraviolet radiation in the Arctic

May 7, 2012, 1:03 pm

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. For a comprehensive review of the extensive and rapidly growing technical literature on this subject, readers are referred to several recent books[4], and particularly to Hessen[5] with its focus on the Arctic. UV-B optics in marine waters and ozone layer depletion and solar ultraviolet radiation are described in Chapter 5. 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[6]. Since ozone layer depletion is expected to continue for many more years, albeit at a slower rate[7], the possible impacts of solar UV-B radiation on marine organisms and ecosystems are currently being investigated[8]. 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[9]. 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[10]. 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[11].

Ultraviolet radiation also appears to affect biogeochemical cycling within the marine environment and in a manner that could affect overall ecosystem productivity and dynamics[12].

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[13]. UV-B-induced mortality in the early life stages, reduced survival and fecundity in females, and changes in sex ratios have all been reported[14]. UV-B-induced damage to the DNA of crustacean zooplankton has also been detected in samples collected up to 20 meters (m) deep[15]. 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[16]. This indicates that Calanus finmarchicus may be sensitive to variation in incident UV-B radiation. Results for the few other species that have been studied are highly variable with some showing strong negative impacts, while others are resistant[17]. 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[18].

The work of Marinaro and Bernard[19], Pommeranz[20], and Hunter et al.[21] provided clear evidence of the detrimental effect of UV-B radiation on the planktonic early life stages of marine fish. Hunter et al.[22], 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[23]. Atlantic cod eggs were negatively affected by exposure to UV-B radiation in very shallow water; 50 cm deep or less[24]. 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[25], while others conclude that they will not[26].

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[27]. 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[28]. 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[29]. 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[30]. Exposure to UV-A (315–400 nm) radiation induced some of the same negative effects on the immune system[31]. Since high cortisol levels induce immunosuppression in fish[32] 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[33]. 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[34] 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[35]. 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)

 

caption Fig. 9.32. Output of a mathematical simulation model[1] illustrating the relative effects of selected variables on UV-induced mortality in Calanus finmarchicus embryos[2]. 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[36]. 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[37]. Arctic phytoplankton appear more susceptible than antarctic species, possibly owing to deeper surface mixed layers in the Arctic[38]. 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[39]. 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[40] and this includes the polyunsaturated fatty acids (PUFAs)[41]. 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[42]. 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[43]. In Atlantic herring, dietary deficiencies of essential fatty acids, in particular docosahexaenoic acid, reduce the number of rods in the eyes[44] and negatively affect feeding at low light levels[45]. Other negative consequences of essential fatty acid deficits have also been reported[46]. 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.[47] 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.[48] 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[49], 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.[50] showed that ozone thickness could in some instances be the single most important determinant of DNA damage in bacterioplankton.

 

caption 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[3].

 

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 populations, 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
9.2. Physical oceanography
    9.2.1. General features
    9.2.2. Sea ice
    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
9.7. Gaps in knowledge and research needs

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Citation

Committee, I. (2012). Effects of changes in ultraviolet radiation in the Arctic. Retrieved from http://www.eoearth.org/view/article/152365

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