Two alternative mechanisms are suggested for nitrite accumulation in the oxygenated oligotrophic water column: (1) excretion by phytoplankton or (2) microbial oxidation of ammonium (nitrification). This study assessed the role of these 2 mechanisms, based on seasonal and high-resolution diurnal depth profiles of the dissolved inorganic nitrogen (DIN) species (nitrite, ammonium, nitrate) and chlorophyll a in the Gulf of Aqaba, Red Sea. Both mechanisms operated in the water column, but in different seasons; nitrification was the prime process responsible for nitrite accumulation during the stratified summer season and phytoplankton nitrite excretion operated during winter mixing. At the onset of summer stratification two N peaks developed below the photic zone, an ammonium maximum (AM) and below it the primary nitrite maximum (PNM). Both peaks were located at a depth range where phytoplankton are thought to be inactive and not excreting nitrite. During summer stratification, the water column deep chlorophyll maximum (DCM), AM, PNM and the nitracline were ordered by a downward increase in N oxidation state similar to the temporal order of the N-species during nitrification. This similarity, together with the diurnal stability of the PNM and its co-existence with oscillating chlorophyll profiles above the DCM, is consistent with nitrification as the key process forming the PNM. We suggest that transport and reaction control the vertical order and separation of N-species in the water column. The ratios between the rate constants for ammonification, ammonium oxidation, nitrite oxidation and nitrate assimilation were estimated by a simple box model to be 1:3:1.5:0.15, respectively. These field estimates are similar to the ratios between the rate constants measured in laboratory experiments.
, and photosynthetic rates concomitantly doubled from 15 to 35 µg C l -1 d -1 . Water transparency declined, as indicated by the vertical diffusion attenuation coefficient K d for photosynthetically active radiation (PAR), which increased from 0.076 to 0.090 m -1 and decreased the euphotic depth from 60 to 45 m. During this time, a significant increase in silica deposition by the diatoms was also detected. We attribute the mentioned changes in environmental characteristics to wind-generated surface currents. Strong winds (up to 10 m s -1 ) during the measurements enriched the surface layers with unusually high nutrient concentrations within <1 d. Hence, primary production rates were observed at a relatively eutrophic nearshore station (MP) and pelagic station (A 1 , 10 km towards the center of the Gulf). They were compared with rates measured on the subsequent day. Values measured were twice as high as those at the pelagic station during the previous calm day. Routine monitoring programs with monthly or semi-weekly sampling are thus likely to miss brief but significant injections of nutrients, leading to the underestimation of seasonal and annual primary production. Our results demonstrate the impacts of transient events on the function and annual yield of aquatic ecosystems.
Variations in transparent exopolymer particles (TEP), bacterial biomass production (BP) and primary productivity (PP) were followed over 52 h at a deep water station in the Gulf of Aqaba (Eilat, Israel) during the spring, in April 2008. About 20 h after the start of the study, there was a short (~15 h) but intense storm event that probably caused a nutrient pulse and, subsequently, a brief outgrowth of diatoms in the euphotic layer. Concentrations of TEP and BP ranged from 23 to 228 µg gum xanthan equivalents l -1 and from 0.2 to 0.6 µg C l -1 h -1, respectively. Concentrations of TEP and BP were measured in unfiltered and in GF/C (1.2 µm)-prefiltered samples. Most of the TEP (59 ± 21% of total TEP, mean ± SD) were in the smaller (GF/C-filtered) size fraction (0.4-1.2 µm); however, after the crash of the diatom bloom, the majority of TEP were in the >1.2 µm size fraction. In the GF/Cfiltered fraction, BP averaged 59 ± 12% and 93 ± 5% of total BP in the upper water column and from 300 m, respectively. Significant correlations were observed between TEP and BP, suggesting that active heterotrophic bacteria may have been associated with these particles. During the 3 d of our study, PP and BP in the euphotic zone averaged 480 and 225 mg C m -2 d -1 , respectively, suggesting that about half or more of the primary produced carbon was metabolized by heterotrophic bacteria in the upper water column. Coincident with strong mixing caused by the storm, TEP concentrations decreased in the surface water and increased at depth. We suggest that TEP acted to link carbon flux between the primary producers and heterotrophic bacteria, and that the downward movement of TEP from the upper water layers may be an important process in transferring organic carbon to deeper waters of the Gulf of Aquaba. Sinking TEP could provide not only organic carbon substrates for associated bacteria but also form 'hot spots' of elevated microbial metabolism and nutrient cycling throughout the water column.
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