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The production and composition of extracellular polymeric substances (EPS) in axenic batch cultures of the benthic marine epipelic diatoms Navicula salinarum and Cylindrotheca closterium were investigated. EPS was secreted into the medium and the bulk was loosely associated with the cells. Neither N. salinarum nor C. closterium formed a well-defined polysaccharide capsule. EPS of both N. salinarum and C. closterium consisted predominantly of polysaccharide but small quantities of protein were present as well. EPS also contained uronic acids and SO % #− groups. Analysis of monosaccharides using gas chromatography showed that for both species glucose and xylose were the main constituents, but several other monosaccharides were present in smaller quantities. Two fractions of EPS were distinguished : a small amount was secreted into the medium and a second fraction was extracted in water at 30 mC. For both species the two fractions differed somewhat in composition, indicating that they represented two different types of EPS. The EPS produced by N. salinarum and by C. closterium differed in their composition. The rate of EPS production in batch culture was highest during the transition from exponential growth to stationary growth. Negatively charged groups such as uronic acids and sulphated sugars determine the adhesion capacity of EPS and probably play an important role in the stabilization of intertidal sediments on which these diatoms grow and produce biofilms.

The secretion of exopolysaccharide in an axenic culture of the marine benthic diatom Cylindrotheca clostenum was investigated. The central question of the experiments was if polysaccharide secretion was dependent on light and photosynthesis. Cells were incubated in the light, in the dark, or in the light with addition of the inhibitor of Photosystem II,3-(3,4-dichloropheny1)-l, l-dimethyl urea (DCMU). These treatments were also applied to a population of benthic diatoms on an intertidal mudflat in the Westerschelde (Scheldt estuary, The Netherlands). In the light (60 p 0 1 photons m-' S-') C. clostenum showed high rates of polysaccharide secretion, while no secretion was observed in the dark or in the presence of DCMU. No intracellular carbohydrate was converted to exopolysaccharide in the dark or in the light with DCMU added. This indicated that secretion of exopolysaccharide was dependent on oxygenic photosynthesis. Similarly, high rates of exopolysaccharide accumulation were observed during daytime emersion on the mudflat, but not in darkened or DCMU-treated sediment. This demonstrated that the pattern observed in cultures of C. closterium was reproducible in situ. It was observed that dunng daytime emersion patterns of vertical migration in the dark and DCMU-treated plots did not differ from those in the light. This implies that motility was not the steering factor for the observed accumulation of exopolysaccharide in the light. When an axenic culture of C. closterium was incubated under an alternating 12 h light:12 h dark cycle, exopolysaccharide concentrations decreased in the dark. Degradation of exopolysaccharide was also observed in the natural population on the mudflat during emersion at night. Because no bacteria were present in the C. clostenum cultures, it was conceived that the degradation of exopolysaccharide observed in cultures was due to secretion of hydrolytic enzymes by C. clostenum.

Sechment bacterial production, abundance and P-glucosidase activity were studied in relation to carbohydrate concentrations in diatom-inhabited intertidal mudflats during the tidal cycle in the Ems-Dollard estuary in summer 1996 During the diurnally emerged period significant increases in water-and EDTA-extracted carbohydrates, bacterial production and abundance in the sediment surface layer (0 to 2 mm) were established. Exposure had no effect on algal biomass (chlorophyll a), doubling times of bacteria or P-glucosidase activity in the surface layer. Net changes in water-extracted carbohydrates together with net changes in total and CTC (5-cyano-2,3-ditolyl tetrazolium chloride)-stained bacteria accounted for 85% of the variations in net increase in bacterial production during the emerged period. During the submerged period significant decreases of water-extracted carbohydrate concentrations occurred. The changes in P-glucosidase activity and water-extracted carbohydrates could explain 93% of the net changes in bacterial production during the submerged period. Waterextracted carbohydrates derived mainly from live diatoms. The source of EDTA-extracted carbohydrates was less specific, but related to chlorophyll a and pheopigments. The results suggest that the net increases in water-extracted carbohydrates due to &atom activities are inbrectly coupled to benthic heterotrophic bacterial production. Bacteria do respond to tidal variations in organic compounds, and water-extracted carbohydrates are an important indicator.

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