Fjord sediments on the west coast of the arctic archipelago Svalbard were surveyed to understand whether large filamentous sulfur bacteria of the genus Beggiatoa thrive at seawater temperatures permanently near freezing. Two sediments had abundant populations of Beggiatoa, while at six sites, only sporadic occurrences were observed. We conclude that Beggiatoa, although previously unnoticed, are widespread in these arctic fjord sediments. Beggiatoa ranged in diameter from 2 to 52 mm and, by those tested, stored nitrate in vacuoles at up to 260 mM. The 16S rRNA gene sequence of a 20-mm-wide filament is closely associated with other large, marine, nitrate-storing Beggiatoa. The Beggiatoa mostly occurred in the upper 2-5 cm of oxidized surface sediment between oxygen and the deeper sulfidic zone. In spite of a very low or an undetectable sulfide concentration, sulfate reduction provided abundant H 2 S in this zone. The total living biomass of Beggiatoa filaments at one study site varied over 3 years between 1.13 and 3.36 g m À2 . Because of their large size, Beggiatoa accounted for up to 15% of the total prokaryotic biomass, even though the filament counts at this site were rather low, comprising o 1/10 000 of the bacterial numbers on a cell basis.

The response of gliding motility to changing temperatures was studied in filaments of the large sulfur bacteria Beggiatoa from arctic, temperate and tropical marine environments. The general shape of the gliding speed vs. temperature curves from all three locations was similar, but differed in the maximal gliding speed of the filaments, optimum temperature and the temperature range of motility. The optimum temperature and the overall temperature range of gliding motility accorded to the climatic origin of the filaments with a high temperature range for tropical, an intermediate range for temperate, and a low temperature range for arctic filaments. The temperature-controlled decrease in gliding speed at low temperatures was reversible while the decline in speed at high temperatures was due to irreversible thermal damage in individual filaments. Filaments from the Arctic and cold-acclimatized filaments from the temperate zone were unaffected by transient freezing of the surrounding seawater. At in situ temperatures, filaments glided at 17-55% of the gliding speed at the optimum temperatures, indicating that they were well adapted to the temperature regime of their origin. Our results point towards an enzymatic control of temperature-dependent gliding motility.

The large sulfur bacteria, Beggiatoa spp., live on the oxidation of sulfide with oxygen or nitrate, but avoid high concentrations of both sulfide and oxygen. As gliding filaments, they rely on reversals in the gliding direction to find their preferred environment, the oxygen–sulfide interface. We observed the chemotactic patterns of single filaments in a transparent agar medium and scored their reversals and the glided distances between reversals. Filaments within the preferred microenvironment glided distances shorter than their own length between reversals that anchored them in their position as a microbial mat. Filaments in the oxic region above the mat or in the sulfidic, anoxic region below the mat glided distances longer than the filament length between reversals. This reversal behavior resulted in a diffusion‐like spreading of the filaments. A numerical model of such gliding filaments was constructed based on our observations. The model was applied to virtual filaments in the oxygen‐ and sulfide‐free zone of the sediment, which is a main habitat of Beggiatoa in the natural environment. The model predicts a long residence time of the virtual filament in the suboxic zone and explains why Beggiatoa accumulate high nitrate concentrations in internal vacuoles as an alternative electron acceptor to oxygen.