Active expression of putative ammonia monooxygenase gene subunit A (amoA) of marine group I Crenarchaeota has been detected in the Black Sea water column. It reached its maximum, as quantified by reverse-transcription quantitative PCR, exactly at the nitrate maximum or the nitrification zone modeled in the lower oxic zone. Crenarchaeal amoA expression could explain 74.5% of the nitrite variations in the lower oxic zone. In comparison, amoA expression by ␥-proteobacterial ammonia-oxidizing bacteria (AOB) showed two distinct maxima, one in the modeled nitrification zone and one in the suboxic zone. Neither the amoA expression by crenarchaea nor that by ␤-proteobacterial AOB was significantly elevated in this latter zone. Nitrification in the suboxic zone, most likely microaerobic in nature, was verified by 15 ammonia-oxidizing bacteria ͉ amoA gene expression ͉ marine group ͉ Crenarchaeota ͉ marine nitrogen loss N itrification, the stepwise oxidation of ammonium to nitrite and then nitrate, is a key process in marine nitrogen cycling. It is responsible for the formation of the large deep-sea nitrate reservoir. It connects the recycling of organic nitrogen to the ultimate nitrogen loss from the oceans, because its products are substrates for denitrification and anaerobic ammonium oxidation (anammox), the only two presently known nitrogen loss processes. In productive waters such as upwelling regions, high fluxes of organic matter and thus remineralization create strong subsurface oxygen minima, enabling denitrification (1-4) or anammox (5-8) to occur. Nitrogen losses from these oxygen minimum zones (OMZs) are estimated to account for 30-50% of total nitrogen loss from the oceans (9, 10). Because remineralization also releases large amounts of ammonium, high nitrification rates are often associated with these OMZs (11), implying that nitrification may play an important role in promoting marine nitrogen loss.The Black Sea is the largest marine anoxic basin in the world. A 20-to 40-m-thick suboxic transitional zone, characterized by low oxygen (Ͻ5 M) and undetectable sulfide, persists throughout the basin between the surface oxic layer and the sulfidic anoxic deep water (Ն100 m) (12, 13). The exact depth zonation varies according to the location within the basin because of circulation and gyre formation, but similar concentrations of chemical species can be traced along isopycnals or density ( t ) surfaces throughout the basin (12). Therefore, the Black Sea provides an ideal model system to study nitrogen cycling processes along oxygen gradients. Nitrification has been reported in the lower oxic zone (14) and so has nitrogen loss via anammox in the suboxic zone (15). Nevertheless, the identity and abundance of the responsible nitrifiers, or any coupling between nitrification and nitrogen losses, remain poorly documented.The first and rate-limiting step of nitrification is aerobic ammonia oxidation. It is a microbially mediated reaction. For decades, only specific groups of ␤-and ␥-proteobacteria have been found to exh...

The oxygen-consuming processes in the hypolimnia of freshwater lakes leading to deep-water anoxia are still not well understood, thereby constraining suitable management concepts. This study presents data obtained from 11 eutrophic lakes and suggests a model describing the consumption of dissolved oxygen (O(2)) in the hypolimnia of eutrophic lakes as a result of only two fundamental processes: O(2) is consumed (i) by settled organic material at the sediment surface and (ii) by reduced substances diffusing from the sediment. Apart from a lake's productivity, its benthic O(2) consumption depends on the O(2) concentration in the water overlying the sediment and the molecular O(2) diffusion to the sediment. On the basis of observational evidence of long-term monitoring data from 11 eutrophic lakes, we found that the areal hypolimnetic mineralization rate ranging from 0.47 to 1.31 g of O(2) m(-2) d(-1) (average 0.90 ± 0.30) is a function of (i) a benthic flux of reduced substances (0.37 ± 0.12 g of O(2) m(-2) d(-1)) and (ii) an O(2) consumption which linearly increases with the mean hypolimnion thickness (z(H)) up to ~25 m. This model has important implications for predicting and interpreting the response of lakes and reservoirs to restoration measures.

We performed combined in situ measurements of bottom boundary-layer turbulence and of diffusive oxygen fluxes at the sediment-water interface in a medium-sized mesotrophic lake. The turbulence was driven by internal seiching with a period of 18 h. This periodic forcing, a prominent feature of enclosed water bodies, led to distinct deviations of the structure and the dynamics of the bottom boundary layer from the classical law-of-the-wall theory. A major feature was a phase lag between the current velocity and the turbulent energy dissipation of approximately 10% of the seiching period (1.5-2 h). The oxygen flux into the sediment was controlled by the diffusive boundary layer, the thickness of which varied between 0.16 and 0.84 mm during the course of a seiching period, and was strongly affected by the periodic bottom boundary-layer turbulence. The rate of dissipation of turbulent energy in the bottom boundary layer allowed us to define the Batchelor length for dissolved oxygen, which quantifies the smallest scales of oxygen fluctuations and provides an appropriate scaling for the diffusive boundary-layer thickness and the corresponding oxygen fluxes. An analysis of the governing time scales revealed the importance of turbulence in controlling the small-scale spatial heterogeneity of the diffusive fluxes. Higher turbulence causes the diffusive boundary layer (DBL) to follow the sediment topography more smoothly, resulting in an increased area-averaged flux due to the greater effective surface area.After surface zones, the bottom boundary layer (BBL) is the second prime site for animals, plants, and microorganisms in natural waters. From a physical and geochemical point of view, the importance of the BBL is twofold. First, the BBL is a major energy sink for basin-scale currents due to bottom friction and also due to the breaking of propagating internal waves on sloping bottoms (Imberger 1998). Consequently, the level of turbulence is enhanced in the BBL compared with the interior water body. Second, the BBL controls the exchange of solutes and particles between water and sediment. The sediment surface is usually an enormous sink of oxygen due to the processes caused by the decomposition of organic matter. Furthermore, the redissolution and subsequent vertical transport of ions and other solutes supply primary producers with nutrients and affect the stability of the water column by chemical (salinity) stratification (Wüest and Gloor 1998).Especially in eutrophic and mesotrophic systems with 1 Corresponding author (andreas.lorke@eawag.ch). AcknowledgmentsWe thank C. Dinkel and M. Schurter for their great help in the field. D. McGinnis kindly improved the English. We gratefully acknowledge the helpful criticism of two unknown reviewers.

Lake Ohrid in southeastern Europe is one of the few ancient, long-lived lakes of the world, and contains more than 200 endemic species. On the basis of integrated monitoring of internal and external nutrient fluxes, a progressing eutrophication was detected (,3.5-fold increase in phosphorus (P) concentration in the lake over the past century). The lake is fortunately still oligotrophic, with high concentrations of dissolved oxygen (DO) in the deep water that are requisite for the unique endemic bottom fauna. Hypolimnetic DO is not only very sensitive to changes in anthropogenic P load-via mineralization of organic material-but also to global warming via decrease of vertical mixing and less frequent complete deep convection. Moreover, these two human effects amplify each other. To keep DO from falling below currently observed minimal levels-given the predicted atmospheric warming of 0.04uC yr 21 -the P load must be decreased by 50% in coming decades. However, even with such a reduction in P load, anoxia is still expected toward the end of the century if the rate of warming follows predictions.

Lake Prespa and Lake Ohrid, located in south-eastern Europe, are two lakes of extraordinary ecological value. Although the upstream Lake Prespa has no surface outflow, its waters reach the 160 m lower Lake Ohrid through underground hydraulic connections. Substantial conservation efforts concentrate on oligotrophic downstream Lake Ohrid, which is famous for its large number of endemic and relict species. In this paper, we present a system analytical approach to assess the role of the mesotrophic upstream Lake Prespa in the ongoing eutrophication of Lake Ohrid. Almost the entire outflow from Lake Prespa is found to flow into Lake Ohrid through karst channels. However, 65% of the transported phosphorus is retained within the aquifer. Thanks to this natural filter, Lake Prespa does not pose an immediate threat to Lake Ohrid. However, a potential future four-fold increase of the current phosphorus load from Lake Prespa would lead to a 20% increase (+0.9 mg P m )3 ) in the current phosphorus content of Lake Ohrid, which could jeopardize its fragile ecosystem. While being a potential future danger to Lake Ohrid, Lake Prespa itself is substantially endangered by water losses to irrigation, which have been shown to amplify its eutrophication.