Increases in ultraviolet radiation at the Earth's surface due to the depletion of the stratospheric ozone layer have recently fuelled interest in the mechanisms of various effects it might have on organisms. DNA is certainly one of the key targets for UV-induced damage in a variety of organisms ranging from bacteria to humans. UV radiation induces two of the most abundant mutagenic and cytotoxic DNA lesions such as cyclobutane–pyrimidine dimers (CPDs) and 6–4 photoproducts (6–4PPs) and their Dewar valence isomers. However, cells have developed a number of repair or tolerance mechanisms to counteract the DNA damage caused by UV or any other stressors. Photoreactivation with the help of the enzyme photolyase is one of the most important and frequently occurring repair mechanisms in a variety of organisms. Excision repair, which can be distinguished into base excision repair (BER) and nucleotide excision repair (NER), also plays an important role in DNA repair
in several organisms with the help of a number of glycosylases and polymerases, respectively. In addition, mechanisms such as mutagenic repair or dimer bypass, recombinational repair, cell-cycle checkpoints, apoptosis and certain alternative repair pathways are also operative in various organisms. This review deals with UV-induced DNA damage and the associated repair mechanisms as well as methods of detecting DNA damage and its future perspectives.

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UV-induced DNA damage and repair: a review

The minute photosynthetic prokaryote Prochlorococcus, which was discovered about 10 years ago, has proven exceptional from several standpoints. Its tiny size (0.5 to 0.7 μm in diameter) makes it the smallest known photosynthetic organism. Its ubiquity within the 40°S to 40°N latitudinal band of oceans and its occurrence at high density from the surface down to depths of 200 m make it presumably the most abundant photosynthetic organism on Earth. Prochlorococcus typically divides once a day in the subsurface layer of oligotrophic areas, where it dominates the photosynthetic biomass. It also possesses a remarkable pigment complement which includes divinyl derivatives of chlorophyll a (Chl a) and Chl b, the so-called Chl a2 and Chl b2, and, in some strains, small amounts of a new type of phycoerythrin. Phylogenetically, Prochlorococcus has also proven fascinating. Recent studies suggest that it evolved from an ancestral cyanobacterium by reducing its cell and genome sizes and by recruiting a protein originally synthesized under conditions of iron depletion to build a reduced antenna system as a replacement for large phycobilisomes. Environmental constraints clearly played a predominant role in Prochlorococcus evolution. Its tiny size is an advantage for its adaptation to nutrient-deprived environments. Furthermore, genetically distinct ecotypes, with different antenna systems and ecophysiological characteristics, are present at depth and in surface waters. This vertical species variation has allowed Prochlorococcus to adapt to the natural light gradient occurring in the upper layer of oceans. The present review critically assesses the basic knowledge acquired about Prochlorococcus both in the ocean and in the laboratory.

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Prochlorococcus, a Marine Photosynthetic Prokaryote of Global Significance

The recent discovery of photosynthetic picoplankton has changed our understanding of marine food webs1. Both prokaryotic2,3 and eukaryotic4,5 species occur in most of the world's oceans and account for a significant proportion of global productivity6. Using shipboard flow cytometry, we have identified a new group of picoplankters which are extremely abundant, and barely visible using traditional microscopic techniques. These cells are smaller than the coccoid cyanobacteria and reach concentrations greater than 105 cells ml–1 in the deep euphotic zone. They fluoresce red and contain a divinyl chlorophyll a-like pigment, as well as chlorophyll b, α-carotene, and zeaxanthin. This unusual combination of pigments, and a distinctive prokaryotic ultrastructure, suggests that these picoplankters are free-living relatives of Prochloron7. They differ from previously reported prochlorophytes—the putative ancestors of the chloroplasts of higher plants—in that they contain α-carotene rather than β-carotene and contain a divinyl chlorophyll a-like pigment as the dominant chlorophyll.

A novel free-living prochlorophyte abundant in the oceanic euphotic zone

Amongst the diverse and potent biological activities of free fatty acids (FFAs) is the ability to kill or inhibit the growth of bacteria. The antibacterial properties of FFAs are used by many organisms to defend against parasitic or pathogenic bacteria. Whilst their antibacterial mode of action is still poorly understood, the prime target of FFA action is the cell membrane, where FFAs disrupt the electron transport chain and oxidative phosphorylation. Besides interfering with cellular energy production, FFA action may also result from the inhibition of enzyme activity, impairment of nutrient uptake, generation of peroxidation and auto-oxidation degradation products or direct lysis of bacterial cells. Their broad spectrum of activity, non-specific mode of action and safety makes them attractive as antibacterial agents for various applications in medicine, agriculture and food preservation, especially where the use of conventional antibiotics is undesirable or prohibited. Moreover, the evolution of inducible FFA-resistant phenotypes is less problematic than with conventional antibiotics. The potential for commercial or biomedical exploitation of antibacterial FFAs, especially for those from natural sources, is discussed.

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Antibacterial free fatty acids: activities, mechanisms of action and biotechnological potential

In the oceans, ubiquitous microscopic phototrophs (phytoplankton) account for approximately half the production of organic matter on Earth. Analyses of satellite-derived phytoplankton concentration (available since 1979) have suggested decadal-scale fluctuations linked to climate forcing, but the length of this record is insufficient to resolve longer-term trends. Here we combine available ocean transparency measurements and in situ chlorophyll observations to estimate the time dependence of phytoplankton biomass at local, regional and global scales since 1899. We observe declines in eight out of ten ocean regions, and estimate a global rate of decline of approximately 1% of the global median per year. Our analyses further reveal interannual to decadal phytoplankton fluctuations superimposed on long-term trends. These fluctuations are strongly correlated with basin-scale climate indices, whereas long-term declining trends are related to increasing sea surface temperatures. We conclude that global phytoplankton concentration has declined over the past century; this decline will need to be considered in future studies of marine ecosystems, geochemical cycling, ocean circulation and fisheries.

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Global phytoplankton decline over the past century

Harmful Marine Algal Blooms

[1] The contribution of ocean-derived DMS to the atmospheric burdens of a variety of sulphur compounds (DMS, MSA, SO2, and nss SO4=) is quantified from season to season. Such quantification, especially for nss SO4= (the climate-relevant product of DMS oxidation), is essential for the quantification of the radiative forcing of climate that may be attributable to marine phytoplankton under possible future climate conditions. Three-dimensional chemical transport modeling up to the stratosphere is used as a tool in realizing this aim. Global data sets on oceanic and terrestrial sulphur sources are used as input. We find that the contribution of ocean-leaving DMS to the global annually averaged column burdens of the modeled compounds is considerable: 11.9 mumol m(-2) (98% of total global burden) for DMS; 0.95 mumol m(-2) (94% of total global burden) for MSA; 2.8 mumol m(-2) (32% of total global burden) for SO2 and 2.5 mumol m(-2) (18% of total global burden) for nss SO4=. The mean annual contribution of DMS to the climate-relevant nss SO4= column burden is greatest in the relatively pristine Southern Hemisphere, where it is estimated at 43%. This contribution is only 9% in the Northern Hemisphere, where anthropogenic sulphur sources are overwhelming. The marine algal-derived input of the other modeled sulphur compounds ( DMS, MSA, and SO2) is also greatest in the Southern Hemisphere where a lower oxidative capacity of the atmosphere, a larger sea-to-air transfer of DMS and a larger emission surface area lead to an elevation of the atmospheric DMS burden.

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The contribution of ocean‐leaving DMS to the global atmospheric burdens of DMS, MSA, SO2, and NSS SO4=

Phytoplankton of the North Sea and its dynamics: A review

We developed a method to investigate the effect of ultraviolet- B radiation (UVBR) on the formation of thymine dimers in microalgal DIVA that can be used for both laboratory and in situ research. Antibody labeling of dimers was followed by a secondary antibody (fluorescein isothiocyanate) staining to allow visualization of DNA damage with flow cytometry or fluorescence microscopy. Thymine dimer-specific fluorescence in nuclear DNA of the marine diatom Cyclotella sp. was linearly related to the UVBR dose. Simultaneous measurements of cellular DNA content showed that the vulnerability of G2 cells to DNA damage did not differ significantly from the vulnerability of G1 cells. The formation and removal of thymine dimers in Cyclotella sp. cells was monitored for 3 consecutive days at two realistic UVBR irradiance bevels. Thymine dimers were removed within 24 h when exposed to a saturating photosynthetically active radiation intensity following the UVBR treatment. This new method allows the study of UVBR- induced DNA damage on a cell-to-cell basis. It is also feasible for field studies because cells remain intact and can be recognized readily after antibody treatment. [KEYWORDS: Bacillariophyceae; cyclotella; dna damage; monoclonal antibody; thymine dimers; uv-b Phytoplankton photosynthesis; radiation; repair; fibroblasts; inhibition; antarctica; dosimeter; growth; impact; ocean]

Monitoring ultraviolet-b-induced dna-damage in individual diatom cells by immunofluorescent thymine dimer detection

During October/November 1983 photosynthetic responses of natural phytoplankton from the Scotia Sea and Bransfield strait to light and temperature were examined in incubators. Both assimilation numbers at saturating light levels and the slopes of the light-limited portions of the photosynthesis versus irradiance curves were smaller than in algae from lower latitudes. However, both parameters increased significantly with rising temperatures. Light-saturated photosynthesis on the average exhibited a Q10-value of ca. 4.2 between-1.5°C and +2°C. Light-limited photosynthesis between-1.5°C and +5°C rose at a rate corresponding to a Q10-value of roughly 2.6. Above +5°C, temperature enhancement of both light-saturated and light-limited photosynthetic rates was minimal or absent. Our results suggest that under extremely low temperatures light-limited photosynthetic rates become temperature-dependent due to changes in maximum quantum yields.

Winfried W.C. Gieskes

Papers

Harmful Marine Algal Blooms

The contribution of ocean‐leaving DMS to the global atmospheric burdens of DMS, MSA, SO2, and NSS SO4=

Phytoplankton of the North Sea and its dynamics: A review

Monitoring ultraviolet-b-induced dna-damage in individual diatom cells by immunofluorescent thymine dimer detection

Light-temperature interactions in the control of photosynthesis in Antarctic phytoplankton