We examined nitrification in the euphotic zone, its impact on the nitrogen cycles, and the controlling factors along a 7500 km transect from the equatorial Pacific Ocean to the Arctic Ocean. Ammonia oxidation occurred in the euphotic zone at most of the stations. The gene and transcript abundances for ammonia oxidation indicated that the shallow clade archaea were the major ammonia oxidizers throughout the study regions. Ammonia oxidation accounted for up to 87.4% (average 55.6%) of the rate of nitrate assimilation in the subtropical oligotrophic region. However, in the shallow Bering and Chukchi sea shelves (bottom ⩽67 m), the percentage was small (0–4.74%) because ammonia oxidation and the abundance of ammonia oxidizers were low, the light environment being one possible explanation for the low activity. With the exception of the shallow bottom stations, depth-integrated ammonia oxidation was positively correlated with depth-integrated primary production. Ammonia oxidation was low in the high-nutrient low-chlorophyll subarctic region and high in the Bering Sea Green Belt, and primary production in both was influenced by micronutrient supply. An ammonium kinetics experiment demonstrated that ammonia oxidation did not increase significantly with the addition of 31–1560 nm ammonium at most stations except in the Bering Sea Green Belt. Thus, the relationship between ammonia oxidation and primary production does not simply indicate that ammonia oxidation increased with ammonium supply through decomposition of organic matter produced by primary production but that ammonia oxidation might also be controlled by micronutrient availability as with primary production.
Nitrogen‐fixing microorganisms (diazotrophs) provide biologically available nitrogen to plankton communities and thereby greatly influence the productivity in many marine regions. Various cyanobacterial groups have traditionally been considered the major oceanic diazotrophs, but later noncyanobacterial and presumably heterotrophic diazotrophs were also found to be widespread and potentially important in nitrogen fixation. However, the distribution and activity of different diazotroph groups is still poorly constrained for most oceanic ecosystems. Here we examined diazotroph community structure and activity along a 7500 km south‐north transect between the central equatorial Pacific and the Bering Sea. Nitrogen fixation contributed up to 84% of new production in the upper waters of the subtropical gyre, where the diazotroph community included the gammaproteobacterium γ‐24774A11 and highly active cyanobacterial phylotypes (>50% of total nifH transcript abundance). Nitrogen fixation was sometimes detectable down to 150 m depth and extended horizontally to the edge of the gyre at around 35°N. Nitrogen fixation was even detected far north on the Bering Sea shelf. In the Alaskan Coastal Waters on the Bering Sea shelf, low nitrate together with high dissolved iron concentrations seemed to foster diazotroph growth, including a prominent role of UCYN‐A2, which was abundant near the surface (1.2×105 nifH gene copies L−1). Our study provides evidence for nitrogen fixation in the Bering Sea and suggests a clear contrast in the composition of diazotrophs between the tropical/subtropical gyre and the separate waters in the cold northern regions of the North Pacific.
In the subtropical oceans, nutrient concentrations are frequently below the detection limits of standard analytical methods. We applied a highly sensitive method to the surface water of the western and central Pacific between 42°N and 40°S and between 141°E and 158°W except in the equatorial zone, and detected overall depletion of nitrate + nitrite and an excess of SRP. However, a remarkable exception was found: an almost complete exhaustion of SRP (<10 nM) existed at a horizontal scale of >2000 km in the western subtropical North Pacific in both summer and winter. The SRP exhaustion was a consequence of an elevated dinitrogen fixation, which occurred in areas with high dust deposition from the Asian continent that likely enhanced SRP consumption. A coupling among nutrient dynamics, dinitrogen fixation and dust deposition produces the extremely low P availability spanning a large area, which appears to be unique to the western North Pacific.
Latitudinal distribution of diazotrophs and their nitrogen (N 2 ) fixation activity were investigated in the western North Pacific in winter (Nov to Dec 2004) and summer (May to Jun 2005) along meridional transects from 37uN to the equator. N 2 fixation activity in whole seawater and seawater passed through a 10-mm filter was assayed by acetylene reduction. The whole-water N 2 fixation was markedly elevated in winter throughout the study area compared to that in summer, probably due to the increased upward supply of phosphate as a result of deeper mixed layer in winter. During both periods a distinct latitudinal variation was observed in N 2 fixation of the whole-water samples at the surface; further, higher activity was observed between the Kuroshio Extension and the salinity front in the North Equatorial Current than in the neighboring areas. The elevated N 2 fixation was primarily ascribed to ,10-mm diazotrophs during both seasons. Flow cytometry conducted in summer revealed that distribution of nanoplanktonic cyanobacteria was closely correlated with that of N 2 fixation activity in the ,10-mm fraction, indicating that nanoplanktonic cyanobacteria were the major diazotrophs in that area. In contrast, microplanktonic diazotrophs, Trichodesmium spp. and Richelia intracellularis exhibited different latitudinal distributions from that of nanoplanktonic cyanobacteria, with maximum numerical abundance of R. intracellularis around 8uN and 30uN, and that of Trichodesmium spp. at 26.5uN. Few microplanktonic diazotrophs occurred in the winter. The distribution of the diazotrophs and their N 2 fixation activity may be controlled by the supply of phosphate and aeolian dust deposition.
Surface ocean phosphate is commonly below the standard analytical detection limits, leading to an incomplete picture of the global variation and biogeochemical role of phosphate. A global compilation of phosphate measured using high-sensitivity methods revealed several previously unrecognized low-phosphate areas and clear regional differences. Both observational climatologies and Earth system models (ESMs) systematically overestimated surface phosphate. Furthermore, ESMs misrepresented the relationships between phosphate, phytoplankton biomass, and primary productivity. Atmospheric iron input and nitrogen fixation are known important controls on surface phosphate, but model simulations showed that differences in the iron-to-macronutrient ratio in the vertical nutrient supply and surface lateral transport are additional drivers of phosphate concentrations. Our study demonstrates the importance of accurately quantifying nutrients for understanding the regulation of ocean ecosystems and biogeochemistry now and under future climate conditions.
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