D.-P. Häder (Germany), H.D. Kumar (India), R.
C. Smith (U.S.A.)
and R. C. Worrest (U.S.A)
Dissolved organic carbon (DOC) and particulate organic carbon (POC) are degradation products of living organisms. These substances are of importance in the cycling of carbon in aquatic ecosystems. UV-B radiation has been found to break down high molecular weight substances and make them available to bacterial degradation. In addition, DOC is responsible for short-wavelength absorption in the water column. Especially in coastal areas and freshwater ecosystems, penetration of solar radiation is limited by high concentrations of dissolved and particulate matter. On the other hand, climate warming and acidification result in faster degradation of these substances and thus enhance the penetration of UV radiation into the water column.
Several research groups have investigated light penetration into the water column. Past studies on UV penetration into the water column were based on temporally and spatially scattered measurements. The process of spectral attenuation of radiant energy in natural waters is well understood and straight forward to model. Less known is the spatial and temporal variability of in-water optical properties influencing UV attenuation and there are few long-term observations. In Europe, this deficiency of measurements is being corrected by a project involving the development of a monitoring system (ELDONET) for solar radiation using three-channel dosimeters (UV-A, UV-B, PAR) that are being installed from Abisko (North Sweden, 68°N, 19°E) to Tenerife (Canary Islands, 27°N, 17°W). Some of the instruments have been installed in the water column (North Sea, Baltic Sea, Kattegat, East and Western Mediterranean, North Atlantic), establishing the first network of underwater dosimeters for continuous monitoring.
Bacteria play a vital role in mineralization of organic matter and provide a trophic link to higher organisms. New techniques have substantially changed our perception of the role of bacteria in aquatic ecosystems over the recent past and bacterioplankton productivity is far greater than previously thought, having high division and turnover rates. It has been shown that bacterioplankton plays a central role in the carbon flux in aquatic ecosystems by taking up dissolved organic carbon (DOC) and remineralizing the carbon. Bacterioplankton are more prone to UV-B stress than larger eukaryotic organisms and, based on one study, produce about double the amount of caeyclobutane dimers.Recently, the mechanism of nitrogen fixation by cyanobacteria has been shown to be affected by UV-B stress. Wetlands constitute important ecosystems both in the tropics and at temperate latitudes. In these areas, cyanobacteria form major constituents in microbial mats. The organisms optimize their position in the community by vertical migration in the mat which is controlled by both visible radiation and UV-B. Cyanobacteria are also important in tropical and subtropical rice paddy fields where they contribute significantly to the availability of nitrogen. Solar UV affects growth, development, and several physiological responses of these organisms.
On a global basis, phytoplankton is the most important biomass producer in aquatic ecosystems. The organisms populate the top layers of the oceans and freshwater habitats where they receive sufficient solar radiation for photosynthetic processes. New research strengthens previous evidence that solar UV affects growth and reproduction, photosynthetic energy harvesting enzymes, and other cellular proteins, as well as photosynthetic pigment contents. The uptake of ammonium and nitrate is affected by solar radiation in phytoplankton, as well as in macroalgae. Damage to phytoplankton at the molecular, cellular, population and community levels has been demonstrated. In contrast, at the ecosystem level there are few convincing data with respect to effects of ozone-related UV-B increases and considerable uncertainty remains. Following UV-B irradiation, shifts in phytoplankton community structure have been demonstrated, which may have consequences for the food web.
Macroalgae and seagrasses are important biomass producers in aquatic ecosystems (but considerably smaller than phytoplankton). In contrast to phytoplankton most of these organisms are sessile and can thus not avoid the exposure to solar radiation at their growth site. Recent investigations showed a pronounced sensitivity to solar UV-B, and effects have been found throughout the top 10-15 m of the water column. Photoinhibition can be quantified by oxygen exchange or by PAM (pulse amplitude modulated) fluorescence. Surface-adapted macroalgae, such as several brown and green algae, show a maximum of oxygen production at or close to the surface; whereas algae adapted to lower irradiances usually thrive best when exposed deeper in the water column. Mechanisms of protection and repair are being investigated.
UV effects on aquatic animals is of increased interest. Evidence for UV effects have been demonstrated in zooplankton activity. Other UV-B sensitive aquatic organisms include sea urchins, corals, and amphibians. Solar UV radiation has been known to affect corals directly. In addition, photosynthesis in their symbiotic algae is impaired, resulting in reduced organic carbon supply. Amphibian populations are in serious decline in many areas of the world, and scientists are seeking explanations for this phenomenon. Most amphibian population declines are probably due to habitat destruction or habitat alteration. Some declines are probably the result of natural population fluctuations. Other explanations for the population declines and reductions in range include disease, pollution, atmospheric changes and introduced competitors and predators. UV-B radiation is one agent that may act in conjunction with other stresses to adversely affect amphibian populations.
The succession of algal communities is controlled by a complex array of external conditions, stress factors and interspecies influences. Freshwater ecosystems have a high turnover and the success of an individual species is difficult to predict but the development of general patterns of community structure follows defined routes. There is a strong predictive relationship between DOC concentration and the depth to which UV radiation penetrates in lakes. Since DOC varies widely, freshwater systems display a wide range of sensitivity to UV penetration. In these systems, increased solar UV-B is an additional stress factor that may change species composition and biomass productivity.
The Arctic aquatic ecosystem is one of the most productive ecosystems on earth and is a source of fish and crustaceans for human consumption. Both endemic and migratory species breed and reproduce in this ocean in spring and early summer, at a time when recorded increases in UV-B radiation are maximal. Productivity in the Arctic ocean has been reported to be higher and more heterogeneous than in the Antarctic ocean. In the Bering Sea, the sea-edge communities contribute about 40–50% of the total productivity. Because of the shallow water and the prominent stratification of the water layer, the phytoplankton is more exposed and affected by solar UV-B radiation. In addition, many economically important fish (e.g., herring, pollock, cod and salmon) spawn in shallow waters where they are exposed to increased solar UV-B radiation. Many of the eggs and early larval stages are found at or near the surface. Consequently, reduced productivity of fish and other marine crops are possible but has not been demonstrated.
There is increased consensus, covering
a wide range of aquatic ecosystems, that environmental UV-B, independent
of ozone related increases, is an important ecological stress that influences
the growth, survival and distribution of phytoplankton. Polar ecosystems,
where ozone-related UV-B increases are the greatest and which are globally
significant ecosystems, are of particular concern. However, these ecosystems
are characterized by large spatial and temporal variability which makes
it difficult to separate out UV-B specific effects on single species or
whole phytoplankton communities. There is clear evidence for short-term
effects. In one study a 4 to 23% photoinhibition of photosystem II activity
was measured under the ozone hole. However, extrapolation of short-term
effects to long-term ecological consequences requires accounting for various
complex effects and quantitative evaluation remains uncertain.