The concentration-dependent uptake of nitrate (NO,) and its inhibit ion by ammonium (NH,) were surveyed in surface waters of the North Atlantic Ocean. Low-level NO, determinations combined with conventional tracer methods provided the first comprehensive data set on NO, utlization kinetics at nanomolar concentrations. Uptake followed saturation kinetics described by the Michaeli s-Menten equation. The half-saturation parameter for uptake (KN) ranged 2-3 orders of magnitude, covarying with ambient NO, concentrations. KN concentrations in oceanic waters averaged -20-30 nM. NH, half-saturation parameters could only be approximated (i.e. KA + A), but observations suggested that KA and K,, were of similar magnitude in oceanic waters. Maximum uptake rates of nitrate, prntN), and ammonium, prncA), covaried, but prncA) almost always exceeded prncN); in oceanic waters, the disparity was an order of magnitude or greater. Most of the variability in A,,(~ and prncA) could be explained by variations in phytoplankton biomass and temperature. The slope of the uptake vs. concentration relationship, CX, was also investigated but was highly variable and could not be related to any of the oceanographic properties observed; cr, was generally greater than ayN. Kinetics analysis showed that NH4 is preferentially utilized over NO3 over the full spectrum of nitrogen concentrations, nanomolar to micromolar.The inhibition of NO3 uptake by NH,, was also parameterized using the Michaelis-Menten expression. The inhibition half-saturation parameter (KJ covaried with KN, but Kj concentrations in oceanic waters (N 40-50 nM) always exceeded KW Maximum inhibition (1,) was rarely complete (i.e. I,,., < I), even at 2,000 nM ammonium. Overall, results suggest that nitrogen utilization parameters currently used in ecosystem models of the open ocean should be re-examined.
Estimates of nutrient demand by dense mats of ice algae in the high Arctic indicate that substantial nutrient fluxes are necessary to satisfy the observed growth over the 2-month bloom. In our study area, Barrow Strait, the quantity of nutrients in the surface-mixed layer is about 3-10 times greater than estimates of total demand during the bloom, and nutrient fluxes in the water column are estimated to be of the same order of magnitude as algal demand. The fluxes in the water column are predicted to vary by more than an order of magnitude over the fortnightly tidal cycle, assuming that fluxes depend upon the strength of tidal currents and the vertical nutrient gradients. In the latter half of the bloom, when biomass levels are high, it appears that established populations of ice algae may experience cyclic conditions of nutrient limitation during neap tides when nutrient fluxes are minimal. Contributions from regeneration and brine exclusion from the ice sheet appear to satisfy only a portion of the bloom's total requirement for nutrients. INTRODUCTIONDense growths of ice algae are a ubiquitous feature beneath annual sea ice in polar regions during the springtime. In the high Arctic these microalgae are largely concentrated at the ice-water interface on or within the skeletal layer of the congelation ice. MacroScopically, they resemble a mottled goldenbrown carpet about 1-2 cm thick. We have found that these algae grow for at least 2 months (April-May) in the central portion of the Northwest Passage. Moreover, the ice alga. e often achieve high levels of biomass (80-100 mg chlorophyll m -e) that are similar to water column values integrated over the depth of the euphotic zone in many planktonic systems.
During mid-May-early June 1997 observations of hydrography, phytoplankton and nitrate concentrations, and abundance and stage distribution of Calanus finmarchicus populations were made in the Labrador Sea and south of Greenland. Egg production rates were also measured for isolated C. finmarchicus females. Surface nitrate and integrated phytoplankton concentrations indicated that, in the deep water, the phytoplankton bloom had ended in the north and east, was in progress in the north central Labrador Sea and near the basin margins, and had not yet become established in an area stretching from the central Labrador Sea to the south of Greenland. C. finmarchicus egg production rates and stage distributions at stations in the 3 areas designated as early, mid-and late/post-bloom zones, suggested that development rates of the overwintered GO generation into mature adults (females and males) were probably low before the bloom, but accelerated during its development. Individual and area1 rates of egg production were highest in the early bloom zone, whereas nauplii were more abundant in the bloom and late/post-bloom zones. Differences in naupliar abundance may have been related to food limitation, or predation. Following development through to the young copepodite stages (CI-111), which were most abundant in the late/post-bloom zone, morality rates were apparently lower and growth rates less dependent on high phytoplankton concentrations and perhaps more dependent on temperature. In the Labrador Sea, where the annual growth season is relatively short and C. finmarchicus produces only 1 generation per year, the timing of the spring bloom may have a significant impact on recruitment of the new year's generation. In areas where the bloom is early and intense, maturation of the overwintered adults will be rapid and egg-laying will occur when phytoplankton concentrations are high. Subsequent survival success of eggs through to later stages will also probably be relatively high and individuals from the new year's generation will have ample time to reach stages capable of overwintering. By contrast, if the bloom is late or of low intensity, adult maturation wdl be delayed and egg-laying may occur when phytoplankton concentrations are low. Under these condtions relatively few eggs may survive and individuals that do survive will have a shorter period in which to attain stages which can overwinter.
The vertical fine structure of particulate matter and inorganic nutrients through the bottom layers of sea ice was determined at a site in the Canadian high arctic. Intense vertical gradients of chlorophyll a, nitrate, ammonium, nitrite, phosphate, and silicate developed in the lower 6 cm of the ice as ice algae attained standing crops of 250 mg∙m−2 (up to 60 mg∙L−1) of chlorophyll a. Pigment and inorganic nutrient concentrations were closely correlated, and pools of inorganic nutrient were shown to exist in the particulate matter, suggesting that the extremely high dissolved nutrient concentrations in the bottom ice (e.g. up to 400 μmol∙L−1 nitrate) were derived at least in part from leakage of algal intracellular pools. Nitrogen and phosphorus were present in excess of algal needs, but silicon may not have been. The ratios of particulate organic carbon to chlorophyll a and to particulate organic nitrogen increased from the ice–water interface upwards, consistent with a physiological response of the ice algae to the strong corresponding gradient of light availability. Although spatially compact, the assemblage of algae in the bottom ice inhabits a highly stratified environment.
Except in "bottom ice" (lowest few centimetres) and surface waters impacted by autotrophs, the major inorganic nutrients behave conservatively in seawater and sea ice. From mid- to late spring, steep and persistent nutrient gradients were observed in the "well-mixed surface layer" with minima near the ice–water interface. Nitrate, ammonium, and phosphate are highly concentrated in heavily colonized bottom ice relative to seawater and the remainder of the ice sheet; concentrations in darkened, weakly colonized bottom ice are similar to the ice sheet. These nutrients also display strong vertical stratification over millimetre scales. Nitrate and phosphate in the bottom ice layer display strong positive relationships with chlorophyll. The accumulation of these nutrients in bottom ice must be biologically mediated and constitutes a significant sink. In contrast, silicic acid concentrations in bottom ice are close to those expected for sea ice formed from the source seawater, are only weakly related to algal biomass, and vary much less seasonally. Ice algae are apparently shocked osmotically and release their intracellular pools of dissolved nutrients. Intracellular pools of nitrate averaged 1.4–9.5% of total particulate nitrogen. Nitrient stresses, during periods of high biomass and sluggish supply, may be alleviated by pooling.
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