Nitrogen utilisation by size‐fractionated phytoplankton assemblages associated with an upwelling event off Westland,New Zealand

Abstract

Uptake rates of ammonium (NH₄ ⁺ ), nitrate (NO₃ ^? ), and urea by three subgroups of phytoplankton (< 200, < 20, < 2 μm) off Westland, were measured using ¹⁵ N tracer techniques in midwinter 1988, after a recent upwelling. For all size fractions at surface irradiance (I₁₀₀), nitrogen (N) was taken up primarily as NO₃ ^?. This accounted for 67–85% of total N uptake (SρN), whereas at 40 and 7% of surface irradiance, the regenerated N (NH₄ ⁺ ) and urea) made up 31–72% of SρN. Depth profile experiments for all three size components showed that uptake of NO₃ ^? was most light‐sensitive, followed by that of NH₄ ⁺ and urea. The irradiance and nutrient availability plot indicated that light was substantially more important than the nutrient concentrations in controlling the assimilation of N by microplankton (20–200 μm). Nano‐ (2–20 μm) and picoplankton (< 2 μm) however, were not as sensitive to either light or nutrient concentrations. High winds and the resulting deep mixing, combined with offshore and alongshore advection in the midwinter, were suggested to be the major cause of the low biomass and N productivity.     

Uptake of nitrogenous nutrients by phytoplankton in a barrier Island estuary: Great South Bay,New York

The uptake of urea, nitrate and ammonium by phytoplankton was measured using ¹⁵N isotopes over a one-year period in Great South Bay, a shallow coastal lagoon. The bay is a unique environment for the study of nutrient uptake since ambient concentrations of $\text{NO}{}_3{}^{\mathrm{?}}\text{NH}{}_4{}^{\mathrm{+}}$ and urea remain relatively high through the year, and phytoplankton are probably never nutrient limited. Urea nitrogen averaged 52% of the total assimilated, while ammonium represented 33% and nitrate 13%. High rates of ammonium uptake occurred only at low urea concentrations (ca< 1-μg-atom urea l^?1). Over the sampling period urea was present in relatively high concentrations, averaging 5·35 μg-atom N l^?1, while means for ammonium and nitrate averaged 1·94 and 0·65 μg-atom N l^?1, respectively. Total N uptake measured with ¹⁵N averaged about 3·3 times the calculated (from elemental ratios and ¹⁴C productivity measurements) N needs of the phytoplankton population. Highest nitrogen uptake occurred in the summer and coincided with the primary production maximum.     

Absorption et régénération de l'azote dans le système brassé de la Manche: productions nouvelle et régénérée

Nitrate and ammonium uptake rates were measured in spring and summer in deep and shallow wellmixed waters of the English Channel during different cruises between 1986 and 1994. In the deep waters, nitrate uptake was relatively low during phytoplankton development and ammonium uptake represented more than 70 % of the total uptake irrespective of the season. In the shallow waters, nitrate uptake during spring phytoplankton growth was high and represented about 75 % of the total uptake. Ammonium uptake became substantial towards the end of spring and summer. The high contribution of ammonium to the nitrogenous nutrition of phytoplankton over the whole of the well-mixed waters is related to a high rate of recycling of nitrogen in the water column. Ammonium regeneration by microheterotrophs can satisfy between 62 % and the totality of the phytoplankton nitrogen requirements. The high primary production in deep well-mixed waters is not supported by allochthonous nitrogen supply but by an intense in situ regeneration of nitrogen. The situation is similar in shallow well-mixed waters, except during the spring bloom.     

Coupling of biological and physical regimes across the Antarctic Polar Front as reflected by nitrogen production and recycling

Oceanographic samples were collected across the Antarctic Polar Front (APF) region in the vicinity of 60°S, 170°W during the US JGOFS program from December 1997 to March 1998. This paper reports the uptake rates of new (nitrate) and regenerated (ammonium and urea) nitrogen measured by ¹⁵N tracer techniques together with the levels of ammonium, urea-N and dissolved free amino acids (DFAAs) during December and mid-February–March. The APF was an important biological boundary, and in December rates of new (nitrate) uptake were greatest south of the APF, exceeding 10 mmol m⁻² d⁻¹ near the retreating ice edge. In February, nitrate uptake rates were an order of magnitude lower. Rates of ammonium uptake in both periods were greater in the warmer water north of the front. Nitrogen f-ratios varied from 0.50 to less than 0.05, with larger values associated with the >5 μm size fraction at the ice edge and generally lower values north of the APF. Urea was an important nitrogen source north of the APF, and lowered f-ratios there by 22% on average when included as part of total nitrogen uptake. Urea uptake was less important south of the APF. Ammonium concentrations increased dramatically south of the APF later in the season, suggesting a system dominated by regeneration. Seasonal changes in the concentrations of regenerated organic compounds such as urea and DFAAs were less obvious, although DFAAs exhibited consistent maxima in the high flow regions of the APF. A mass balance based of ammonium fluxes suggests that nitrification was significant at locations south of the APF in February. In these nitrate-replete waters, light/mixing conditions in the surface water (the Sverdrup criterion) accounted for over 50% of the variance in nitrate uptake rates. The stability responsible for higher new production south of the APF is due both to the separation of this region from the maximum zonal wind field to the north as well as to melt-water contribution from the retreating ice field. Estimated new production and exportable carbon production exceeded 500 mmol nitrate m⁻² yr⁻¹ and 40 g C m⁻² yr⁻¹, respectively, south of the APF. Thus, new production in the marginal ice zone of the Southern Ocean rivals that in coastal systems and indicates that this is an important region for export production.     

海洋浮游生物氮吸收动力学及其粒级特征

于１９９１年秋－１９９２年夏在中国科学院生态网络站之一的胶州湾进行了４个季节的现场实验，运用＾１５Ｎ同位素示踪方法研究胶州湾浮游生物群落对两种主要源铵态氮和硝态氮的吸收动力学及其粒级特征。研究初步阐明自然浮游生物落在不同季节，对不同氮源的吸收特性和受控机制，首次在群落水平上给出不同粒级浮游生物氮吸收特征的定量描述，从而为新生产力研究，生态系能流分配和生源要素生物地化循环研究提供重要参数。     

Studies of nitrogen assimilation by marine phytoplankton and its implication in the nitrogen cycling in the sea

I present here a review of my work concerning nitrogen assimilation by marine phytoplankton. This opportunity was provided to me as the recipient of the Okada Prize for 1990 from the Oceanographical Society of Japan. Assimilation of nitrogenous nutrients by phytoplankton has received considerable research effort since it is an essential process in organic matter production in the sea surface. The use of¹⁵N technique is necessary for tracing nitrogen assimilation by natural marine phytoplankton, but nitrogen metabolism of heterogenous natural populations significantly complicates flow of isotope. Dilution of¹⁵N isotope by heterotrophic regeneration of ammonium causes underestimates of uptake rates. I made an evaluation of isotope dilution effects in available data sets of¹⁵N-ammonium uptake experiments in literature. Incorporated¹⁵N in particulates might revert back to dissolved organic or inorganic nitrogen. I conducted pulse-chase experiments which can quantify such loss of tracer. From these studies, a short term experiment with sufficient amount of tracer enrichment is found to overwhelm these problems. In such an experiment, however, the elevation of nutrient concentration by tracer addition may likely perturb the uptake process. An initial rapid uptake is expected if the population is nitrogen deficient, but I found that this phenomenon is not common to surface oligotrophic open oceans. Uptake rate from such an experiment, or capacity of nitrogen uptake, was obtained using surface waters from an extended area in the North Pacific, and its regional variability was discussed. In addition to overall¹⁵N uptake, time series analysis of intracellular¹⁵N partitioning between hot ethanol soluble and insoluble fractions was found to be useful. When¹⁵N-ammonium is added to nitrogen deficient cells of phytoplankton,¹⁵N is accumulated in the ethanol soluble fraction. Using cultured strains of marine phytoplankton, this accumulation was proved to be caused by the difference of rates of nitrogen uptake and nitrogenous macromolecule synthesis. Uptake rate per cell is relatively constant irrespective of nutritional status, but macromolecule synthesis decreases with nitrogen deficiency. This accumulation of¹⁵N in the ethanol soluble fraction was used as an index of nutritional status with respect to nitrogen of the natural populations of phytoplankton from the western North Pacific. The uptake capacity of nitrate was observed to be higher than that of ammonium in the regional upwelling around Izu Islands and during the spring bloom in Alaskan coastal water. The¹⁵N partitioning technique revealed that nitrate taken up was rapidly incorporated in the macromolecule fraction. This suggests that ammonium uptake is suppressed to be smaller than intracellular nitrogen assimilation, rather than that nitrate is taken up in excess and accumulates within the cell. Regulation of nitrate uptake by light intensity was also discussed in detail for the Alaskan data. Several other studies currently conducted are also mentioned.     

Wintertime rates of net primary production and nitrate and ammonium uptake in the southern Benguela upwelling system

The elevated levels of primary productivity associated with eastern boundary currents are driven by nutrient- rich waters upwelled from depth, such that these regions are typically characterised by high rates of nitrate-fuelled phytoplankton growth. Production studies from the southern Benguela upwelling system (SBUS) tend to be biased towards the summer upwelling season, yet winter data are required to compute annual budgets and understand seasonal variability. Net primary production (NPP) and nitrate and ammonium uptake were measured concurrently at six stations in the SBUS in early winter. While euphotic zone NPP was highest at the stations nearest to the coast and declined with distance from the shore, a greater proportion was potentially exportable from open-ocean surface waters, as indicated by the higher specific nitrate uptake rates and f-ratios (ratio of nitrate uptake to total nitrogen consumption) at the stations located off the continental shelf. Near the coast, phytoplankton growth was predominantly supported by ammonium despite the high ambient nitrate concentrations. Along with ammonium concentrations as high as 3.6 µmol l^–1, this strongly suggests that nitrate uptake in the inshore SBUS, and by extension carbon drawdown, is inhibited by ammonium, at least in winter, although this has also been hypothesised for the summer.     

Primary production and nutrient assimilation in the Iberian upwelling in August 1998

Primary production was measured during two Lagrangian experiments in the Iberian upwelling. The first experiment, in a body of upwelled water, measured day-to-day changes in phytoplankton activity as the water mass moved south along the shelf break. Nutrient concentrations decreased over a five day period, with concomitant increases in phytoplankton biomass. Initially the maximum phytoplankton biomass was in the upper 10m but after four days, a sub-surface chlorophyll maximum was present at 30m. Depth-integrated primary production at the beginning of the experiment was 70mmolC.m⁻².d⁻¹ (838mgC.m⁻².d⁻¹) and reached a maximum of 88mmolC.m⁻².d⁻¹ (1053mgC.m⁻².d⁻¹) on day 3. On day 1, the picoplankton fraction (<2μm) was slightly more productive than larger (>5μm) phytoplankton, but the increase in overall production during the drift experiment was by these larger cells. Nitrate was the dominant nitrogen source. As nutrient concentrations declined, ammonium became increasingly more important as a nitrogen source and the f-ratio decreased from 0.7 to 0.5. Picoplankton cells (<2μm) were responsible for most (65–80%) of the ammonium uptake. The C:N:P uptake ratios were very close to the Redfield ratio for the first four days but as nutrients became depleted high C:N uptake ratios (11 to 43) were measured. Over the period of the experiment, nitrate concentration within the upper 40m decreased by 47.91mmolN.m⁻². In vitro estimates, based on ¹⁵N nitrate uptake, accounted for 56% of the decrease in nitrate concentration observed in the drifting water mass. Ammonium uptake over the same four day period was 16.28mmolN.m⁻², giving a total nitrogen uptake of 43.18mmolN.m⁻².In the second experiment, an offshore filament was the focus and a water mass was sampled as it moved offshore. Nutrient concentrations were very low (nitrate was <10nmol l⁻¹ and ammonium was 20–40nmol l⁻¹). Primary production rate varied between 36mmolC.m⁻².d⁻¹ (436mgC.m⁻².d⁻¹) and 21mmolC.m⁻².d⁻¹ (249mgC.m⁻².d⁻¹). Picophytoplankton was the most productive fraction and was responsible for a constant proportion (ca 0.65) of the total carbon fixation. Uptake rates of both nitrate and ammonium were between 10 and 20% of those measured in the upwelling region. Urea could be a very significant nitrogen source in these waters with much higher uptake rates than nitrate or ammonium; urea turnover times were ca. one day but the source of the urea remains unknown. Urea uptake had a profound effect on calculated f ratios. If only nitrate and ammonium uptake was considered, f ratios were calculated to be 0.42–0.46 but inclusion of urea uptake reduced the f ratio to <0.1. The primary production of this oligotrophic off-shore filament was driven by regenerated nitrogen.     

Seasonal variability in nitrogenous nutrition of phytoplankton assemblages in the northeastern subarctic Pacific Ocean

Nitrogen uptake rates, and physical, chemical and biological characteristics of the euphotic zone were studied in winter, spring and late summer during the period 1992–1994 along a transect (Line P) extending from the continental slope off the southwest corner of Vancouver Island (British Columbia, Canada; station P4; 49°N, 127°W) to open waters in the NE Pacific (OSP; 50°N, 145°W). Nitrate (NO₃⁻) and silicic acid (Si(OH)₄) concentrations increased offshore during every season. Lowest NO₃⁻ and Si(OH)₄ values were observed during late summer and spring, and highest during winter throughout the euphotic zone. For spring and late summer, surface depletion of NO₃⁻ was observed at the inshore end of the transect, while offshore concentrations were never limiting for phytoplankton growth. Silicic acid was never depleted at any depth or station during the period covered by this study. Ammonium (NH₄⁺) and urea concentrations exhibited a patchy distribution along the transect, with no seasonal variations. Chlorophyll a and particulate nitrogen did not show a consistent longitudinal pattern from year to year. In general, the highest concentrations of chlorophyll a and particulate nitrogen were measured during the late summer cruises, with lower values in spring and lowest in winter. Phytoplankton assemblages were numerically dominated by flagellates <5 μm throughout the water column on each cruise transect. Ammonium, urea and NO₃⁻ uptake rates represented on average 55, 24 and 21% of the depth-integrated total nitrogen uptake, both longitudinally and seasonally; hence, phytoplankton utilized nitrogen in the following order: NH₄⁺>urea>NO₃⁻ along Line P. Ammonium may have inhibited the uptake rates of NO₃⁻ and urea. Urea uptake rates were lower than those of NH₄⁺, but higher values were occasionally observed at a few depths along the transect, particularly during the spring of 1993. Depth-integrated NH₄⁺ uptake rates were generally higher inshore, while NO₃⁻ uptake rates showed higher values offshore during most seasons. In contrast, urea uptake rates did not exhibit a consistent longitudinal trend. The depth-integrated f-ratio ranged from 0.05 to 0.37 with an average of 0.21 for all stations and cruises, and was overestimated on average by 36% when urea was excluded from the calculation. On a yearly basis, primary productivity in the NE subarctic Pacific was based on regenerated nitrogen.     

Nitrogen cycling in the southern North Sea: consequences for total nitrogen transport

Nitrate and ammonium uptake rates were measured during a series of cruises in the well-mixed region of the southern North Sea from February to September. Water column-integrated uptake rates ranged between 0.01 and 8.7 mmol N m⁻² d⁻¹ and 0.01 and 12.2 mmol N m⁻² d⁻¹ for nitrate and ammonium, respectively, with ammonium uptake dominating after the phytoplankton spring bloom in May. A moored buoy continuously measuring nitrate and chlorophyll a and seabed current meters were also deployed in the central southern North Sea in the region of the East Anglian plume—a permanent physical feature which transports nutrients towards continental Europe. This enabled the flux of water and hence of nutrients across the southern North Sea to be determined and an assessment of the contribution of freshwater nutrients to the flux to be made. A simple box model is developed to relate the phytoplankton uptake of nitrate and ammonium to the transport of nitrate, ammonium and particulate organic matter (POM) across the southern North Sea. This showed the importance of the plume region of the North Sea in the processing of nitrogen, with nitrate dominating total nitrogen transport prior to the spring bloom (10 340×10³ kg N inflow to the plume in March) and transport of nitrogen as ammonium, nitrate and POM in approximately equivalent amounts during summer (2560, 2960 and 2151×10³ kg N inflow to the plume, respectively, in July). The box model also demonstrates more generally the need to assess nitrogen transport as nitrate, ammonium and POM if an improved understanding of the impact of nutrient input in shelf seas is to be achieved.     

小分子有机碳、氮源对海洋着色菌(Marichromatium gracile)生长和去除高浓度无机三态氮的影响

采用高浓度无机三态氮(铵氮4NH?-N、亚硝氮2NO?-N和硝氮3NO?-N)共存的模拟海水体系,在最适生长条件下,研究了小分子有机物(糖类、有机酸、醇、有机氮)和p H对海洋着色菌(Marichromatium gracile)YL28去除水体无机三态氮的影响。结果表明:以葡萄糖、乙酸钠和乙醇为唯一碳源时,水体中的高浓度2NO?-N和3NO?-N均能被完全去除,4NH?-N的去除率分别为93.40%、84.55%和66.63%;碳源为乙酸钠时菌体生长最好,体系中添加蛋白胨或尿素,仅4NH?-N的去除效果明显降低。p H值在6.0—9.0时,该菌株对4NH?-N、2NO?-N和3NO?-N均具有去除能力。由此可知:YL28菌株对模拟海水养殖水体中高浓度无机三态氮具有良好的去除能力,高浓度有机氮化物(蛋白胨和尿素)对4NH?-N的去除能力有明显影响,但对2NO?-N和3NO?-N仍保持高效的去除能力。本研究为不产氧光合细菌制剂在水产养殖中的合理应用提供参考。     

Ammonium and phosphorus flux through the microplankton community in Agulhas Bank waters

The size-related activities of important heterotrophs and autotrophs were compared at the surface and at the subsurface chlorophyll maximum (Chl_max) in Agulhas Bank waters. The netplankton fraction was dominated by Nitzschia spp. and ciliates, small (diameter c. 3μm) microflagellates being the most abundant nanoplankton group. Uptake ratios of ammonium to phosphate for the total microplankton community were different at the two depths. With reference to the Redfield ratio, it appears that at least 48 per cent of the nitrogen ration at the Chl_max was regenerated even though nitrate was in ample supply. These assimilation ratios also suggest a very large contribution from recycled nitrogen other than ammonium in surface waters. It is unlikely that phosphorus would ever become limiting, except perhaps at the primary production maximum. Microplankton uptake and regeneration of both ammonium and phosphorus were approximately in balance, indicating that variations in assimilation ratios were the result of heterotrophic excretory activity. The size-fractionation studies show that picoplankton were on average the single most important size class in nutrient assimilation. The netplankton size class was, in terms of regeneration, often the most active in the microplankton community especially within the Chl_max. Heterotrophic microflagellates and picoplankton supplied the bulk of ammonium and phosphorus at the surface. The importance of a particular size class to either ammonium or phosphorus uptake/excretion was quantified as a relative assimilation/regeneration index. These calculations demonstrate size-related differences in the relative importance of the microplanktonic groups to the immobilization and recycling of different nutrients.     

Physiology of Laminaria

Abstract. Dissolved nitrate, nitrite, ammonium and phosphate were monitored for 2.5 years along a coastal eutrophication gradient originating at the St. Andrews sewer outfall. Linear growth rates and tissue composition (phosphorus, various nitrogen fractions and storage carbohydrates) were monitored at the same stations in lamina tissue of Laminaria digitata and L. saccharina. Growth rates were considerably enhanced at the eutrophicated stations both in spring, when exogenous nutrients were at peak values at all stations, and during the summer when exogenous nutrients were very low at all stations. Enhanced summer growth rates were correlated with the increased reserves of N and P accumulated during winter and spring, and particularly with soluble organic nitrogen reserves. Accumulation of storage carbohydrates was inversely correlated with growth rate and tissue N and P reserves, presumably because fixed carbon could be incorporated into new protein and thence new tissue only if internal non-protein N reserves were available.     

Hydrographic conditions during the 2004 SBI process experiments

Western Arctic Shelf-Basin Interactions (SBI) process experiment cruises were conducted during spring and summer in 2002 and 2004. A comparison of the 2004 data with the results from 2002 reveals several similarities but also some distinct differences. Similarities included the following: (1) Dissolved inorganic nitrogen (DIN) (ammonium+nitrate+nitrite) limited phytoplankton growth in both years, suggesting that the fixed-N transport through Bering Strait is a major control on biological productivity. (2) The head of Barrow Canyon was a region of enhanced biological production. (3) Plume-like nutrient maxima and N** minima (a signal of sedimentary denitrification) extending from the shelf into the interior were common except at our easternmost section where the nearshore end of these features intersected the slope. (4) Particularly during summer, oxygen supersaturations were common in or just above the shallow nitracline. (5) Surface waters at our deepest stations were already depleted in nitrate, ammonium and urea during our springtime observations. A major difference between the 2 years was the greater influence of warm, relatively low-nutrient Alaska Coastal Water (ACW) during 2004 entering the region via Bering Strait. This increased inflow of ACW may have reduced photic zone nutrient concentrations. The differences in water temperature and nutrients were most pronounced in the upper ∼100 db, and the increased influence of warm water in 2004 relative to 2002 was most evident in our East Barrow (EB) section. Although the EB data were collected on essentially the same year-days (29 July–4 August 2002 vs. 29 July–6 August 2004), the surface layers were up to 5° warmer in 2004.While the stronger inflow of ACW in 2004 may have reduced the autochthonous nutrient supply, rates of primary production, bacterial production, and particulate organic carbon export were higher in 2004. This conundrum might be explained by differences in the availability of light. Although, springtime ice thicknesses were greater in 2004 than in 2002, snow cover was significantly less and may have more than compensated for the modest differences in ice thickness vis a vis light penetration. In addition, there was a rapid and extensive retreat of the ice cover in summer 2004. Increased light penetration in 2004 may have allowed phytoplankton to increase utilization of nutrients in the shallow nitracline. In addition, more light combined with warmer temperatures could enhance that fraction of primary production supported by nutrient recycling. Enhanced subsurface primary production during summer 2004 is suggested not only by the results of incubation experiments but by more extreme dissolved oxygen supersaturations in the vicinity of the nitracline. We cannot, however, ignore aliasing that might arise from somewhat different station distributions and timing. It is also possible that the rapid ice retreat and warmer temperatures lead to an acceleration in the seasonal progression of biological processes such that the summer 2004 SBI Process Cruise (HLY 04-03) experiment was observing a state that might have existed a few weeks after completion of the 2002 summer cruise (HLY 02-03). Despite these complications, there is little doubt that biological conditions at the ensemble of hydrographic stations occupied in 2004 during the SBI Process Cruises differed significantly from those at the stations occupied in 2002.     

夏季南极普里兹湾氮吸收受融冰调控

本研究将稳定同位素（¹⁵N）与放射性核素（²²⁶Ra）示踪相结合,探讨了2006年夏季南极普里兹湾融冰过程对氮吸收的调控作用。硝酸盐及铵盐的比吸收速率均与浓度呈正相关关系,表明底物浓度效应的存在。开阔洋区具有较高的f比值,而在埃默里冰架附近f比值低得多。f比值与冰融水份额之间存在负相关关系,暗示融冰过程在调控水体氮吸收方面起着重要作用。融冰改变了当地上层水体的层化条件,显著地影响南大洋的生物泵效率及对大气二氧化碳的吸收。本研究为南大洋上层水体碳、氮动力学的调控机制提供了新的认识,且有助于对历史记录的解读及对未来气候变化的预测。     

New and regenerated production in the Almeria-Oran front area,eastern Alboran Sea

Uptake and regeneration of nitrogen in the Almeria-Oran frontal zone (SW Mediterranean) and adjacent (Atlantic and Mediterranean) systems were studied during the Almofront I cruise (JGOFS-France). The frontal zone was characterized by an upsloping of nitracline from about 50 m in the adjacent systems to 25–30 m within. Along with nitrate, ammonium, chlorophyll a and particulate organic nitrogen also were at higher concentrations in the frontal zone than in the adjacent waters.The nitrate uptake rates were significantly higher in the frontal zone (up to 6.4 nmol l⁻¹ h⁻¹) than in the Atlantic and Mediterranean waters (generally <1 nmol l⁻¹ h⁻¹) indicating a significant increase of new production at the front. This increase was related to the upsloping of the nitracline as shown by the significant correlation (p<0.05) between new production and depth of the nitracline. The new production in the Almeria-Oran was much lower than those recorded in other oceanic and coastal fronts. This could be related to the fact that the nitracline did not rise up to the surface and the high concentrations of nitrate were confined to deeper layers where the ambient light intensity was less. Nitrate uptake in the frontal zone was significantly higher, by 1.7–5.8 times (average 4.2), than the calculated diffusive flux of nitrate, suggesting that vertical advection may be an important source of nitrate. New production rates at the front were also significantly higher (3–9 times, average 5.8) than the PON flux to 100 m depth estimated by sediments traps (Journal of Marine Systems 5, 377–389), suggesting a strong decoupling between surface production and downward flux of POM in the frontal zone.The north–south gradient observed with different parameters indicates the presence of a transfrontal secondary circulation. This distribution also suggests that the primary production in the front is initially nitrate-based, with a diatom-herbivore food chain, whereas regenerated production, associated with an intense recycling of organic matter, later becomes progressively important in time and space.     

Physiology of Laminaria

Abstract. Sporophytes of Laminaria digitata and L. saccharina were collected from the shore at different times of year. Intact sporophytes of both species, and discs cut from L. digitata lamina tissue were maintained for several weeks in laboratory cultures under various nitrate, phosphate, temperature and daylength regimes. Substrate-saturated uptake rates of approximately 24 μgN g dry weight^-1 h^-1 from 20 μM nitrate and 8 μgP g dry weight^-1 h^-1 from 10 μM phosphate were more than sufficient to account for internal nitrogen and phosphorus accumulation. Other nitrogen sources - nitrite, ammonium and urea - were also taken up, independently of each other, and supported growth. During the late spring decline of ambient nutrient levels, when growth rates on the shore also decline markedly, enrichment with nitrate (15 μM) and phosphate (3μ) together prevented this decline and also maintained peak photosynthetic rates (net photosynthesis = 10.4 μlO₂ cm^-2 h^-1) which otherwise dropped to 4.5 μO₂ cm^-2 h^-1 over 47 days at ambient nutrient levels (0.5 μM nitrate and phosphate). Slow summer growth rates in June/July were enhanced to greater than spring peak values by combined nitrate (7.5 μM) and phosphate (1.8 μM) enrichment; neither was sufficient alone, although the individual nutrients were accumulated internally. A lesser, but significant enhancement was also achieved in September. In midwinter, nutrient enhancement itself (10.5 μM nitrate and 3.0 μM phosphate) had a small effect on growth rates only if summer water temperature (15°C) was used, but the dramatic effect of an increased photoperiod (7.5 to 17.5 h) was in excess of that expected for the increased radiant energy available. This was found to be due, at least in L. digitaia discs, to the re-establishment of a surface meristoderm in the dormant winter tissue; this was particularly active with high phosphate supply. Growth of the new lamina in January on the shore was much reduced if the old lamina was removed, although the time of initiation of new growth remained unchanged. In the laboratory, the old lamina was found to be a source of nitrogen for new lamina but not of fixed carbon or phosphorus. New lamina photosynthesis, manifest as mannitol accumulation, was directly proportional to phosphate supply up to at least 7.5 μM phosphate. Mannitol accumulation was then suppressed by a nitrate supply above 12 um, presumably by diversion of fixed carbon to other biosynthetic pathways.     

Inorganic nitrogen acquisition by the tropical seagrass Halophila stipulacea

The tropical seagrass Halophila stipulacea is dominant in most regions of the Indo‐Pacific and the Red Sea and was introduced into the Mediterranean Sea after the opening of the Suez canal. The species is considered invasive in the Mediterranean Sea and has been progressively colonizing new areas westward. Growth and photosynthetic responses of H. stipulacea have been described but no information is yet available on the nitrogen nutrition of the species. Here we simultaneously investigated the uptake kinetics of ammonium and nitrate and the internal translocation of incorporated nitrogen in H. stipulacea using ¹⁵N‐labelled substrates across a range of N_i levels (5, 25, 50 and 100 μm ). The ammonium uptake rates exceeded the nitrate uptake rates 100‐fold, revealing a limited capacity of H. stipulacea to use nitrate as an alternative nitrogen source. The uptake rates of ammonium by leaves and roots were comparable up to 100 μm ¹⁵NH₄Cl. At this concentration, the leaf uptake rate was 1.4‐fold higher (6.22 ± 0.70 μmol·g^?1 DW h^?1) than the root uptake rate (4.54 ± 0.28 μmol·g^?1 DW h^?1). The uptake of ammonium followed Michaelis–Menten kinetics, whereas nitrate uptake rates were relatively constant at all nutrient concentrations. The maximum ammonium uptake rate (V_max) and the half‐saturation constant (K_m) of leaves (9.79 μmol·g^?1 DW h^?1 and 57.95 μm , respectively) were slightly higher than that of roots (6.09 μmol·g^?1DW h^?1 and 30.85 μm , respectively), whereas the affinity coefficients (α = V_max/K_m) for ammonium of leaves (0.17) and roots (0.20) were comparable, a characteristic that is unique among seagrass species. No substantial translocation (<2.5%) of ¹⁵N incorporated as ammonium was detected between plant parts, whereas the translocation of ¹⁵N incorporated as nitrate was higher (40–100%). We conclude that the N_i acquisition strategy of H. stipulacea, characterized by a similar uptake capacity and efficiency of leaves and roots, favors the geographical expansion potential of the species into areas with variable water‐sediment N levels throughout the Mediterranean.     

西北冰洋中太平洋入流水营养盐的变化特征

利用1999,2003和2008年夏季(7-9月)三次中国北极科学考察数据资料,分析和讨论太平洋入流水营养盐的分布和楚科奇海关键生物地球化学过程对太平洋水化学性质的改造.结果表明,2003和2008年在白令海峡南部64.3°N纬向断面(BS断面)由于水团性质差异显著,营养盐呈西高、东低的分布趋势.2003年BS断面水柱...