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...