Effects of UV radiation on phytoplankton

Abstract
It is now widely documented that reduced ozone will result in increased levels of ultraviolet (UV) radiation, especially UV‐B (280–320nm), incident at the surface of the earth [Watson, 1988; Anderson et al., 1991; Schoeberl and Hartmann, 1991; Frederick and Alberts, 1991; WMO, 1991; Madronich, 1993; Kerr and McElroy, 1993], and there is considerable and increasing evidence that these higher levels of UV‐B radiation may be detrimental to various forms of marine life in the upper layers of the ocean. With respect to aquatic ecosystems, we also know that this biologically‐ damaging mid‐ultraviolet radiation can penetrate to ecologically‐ significant depths in marine and freshwater systems [Jerlov, 1950; Lenoble, 1956; Smith and Baker, 1979; Smith and Baker, 1980; Smith and Baker, 1981; Kirk et al., 1994]. This knowledge, plus the dramatic decline in stratospheric ozone over the Antarctic continent each spring, now known to be caused by anthropogenically released chemicals [Solomon, 1990; Booth et al., 1994], has resulted in increased UV‐environmental research and a number of summary reports. The United Nations Environmental Program (UNEP) has provided recent updates with respect to the effects of ozone depletion on aquatic ecosystems (Hader, Worrest, Kumar in UNEP 1989, 1991, Hader, Worrest, Kumar and Smith UNEP 1994) and the Scientific Committee on Problems of the Environment (SCOPE) has provided [SCOPE, 1992] a summary of the effects of increased UV radiation on biological systems. SCOPE has also reported [SCOPE, 1993] on the effects of increased UV on the biosphere. In addition, several books have recently been published reviewing various aspects of environmental UV photobiology [Young et al., 1993], UV effects on humans, animals and plants [Tevini, 1993], the biological effects of UV radiation in Antarctica [Weiler and Penhale, 1994], and UV research in freshwater ecosystems [Williamson and Zagarese, 1994]. Several other reviews are relevant [NAS, 1984; Caldwell et al., 1986; Worrest, 1986; NOAA, 1987; Smith, 1989; Smith and Baker, 1989; Voytek, 1990; Häder, 1993; Acevedo and Nolan, 1993; Holm‐Hansen et al., 1993; Vincent and Roy, 1993; Biggs and Joyner, 1994; Williamson and Zagarese, 1994; Karentz, 1994; Cullen and Neale, 1993; Cullen and Neale, 1994]. As Hader et al. have summarized [UNEP, 1989; UNEP, 1991], “UV‐B radiation in aquatic systems: 1) affects adaptive strategies (e.g., motility, orientation); 2) impairs important physiological functions (e.g., photosynthesis and enzymatic reactions); and 3) threatens marine organisms during their developmental stages (e.g., the young of finfish, shrimp larvae, crab larvae)”. Possible consequences to aquatic systems include: reduced biomass production; changes in species composition and biodiversity; and alterations of aquatic ecosystems and biogeochemical cycles associated with the above changes. Within the past four years, our knowledge with respect to the environmental effects of ozone‐related increased levels of UV‐B has increased significantly, and numerous efforts have been directed toward process‐oriented studies of UV responses in plants and animals. Consensus is building toward the view that current levels of UV play a major role as an ecological determinant, influencing both survival and distribution, and are thus deserving of increased study independent of ozone‐related UV‐B increases. This review outlines U.S. research subsequent to 1991 and emphasizes studies concerned with phytoplankton.