Effects of a crude oil and an oil dispersant (Corexit 9527) on populations of the littleneck clam (protothaca staminea)

Field and laboratory experiments were carried out to investigate the effects of Alberta crude oil and an oil dispersant (Corexit 9527) on the larval settlement, survival, siphon activities and behaviour of the littleneck clam (Protothaca staminea). Corexit 9527 was much more toxic than crude oil, and the highest toxicity was obtained when Corexit 9527 was mixed with crude oil. Siphon activities were impaired and abnormal behaviour was exhibited when adult clams were treated with 100 ppm Corexit 9527, 1000 ppm crude oil or a combination of both. Larval settlement was not affected when the substratum was treated with 1000ppm crude oil but was retarded when the substratum was treated with a mixture of 1000 ppm oil and 100 ppm Corexit 9527. Gas chromatograms also showed that the retention time and depth of penetration of hydrocarbons in the substratum was increased when Corexit 9527 was used with crude oil.     

The action of heavy metals on the gametes of the marine mussel, Mytilus edulis (L.)-II. Uptake of copper and zinc and their effect on respiration in the sperm and unfertilized egg

The addition of stock copper and zinc nitrate solutions to filtered seawater (pH 7·91) resulted in a large pH shift which was more pronounced with copper. The pH shift was minimized by increasing the buffering capacity of seawater using 10 mm Tris-Tes pH 7·91. Subsequent experimentation was carried out in both unbuffered and buffered seawater.In unbuffered seawater, both copper and zinc had a pronounced inhibitory effect on Mytilus edulis sperm respiration with 50% inhibition occurring at 0.65 mm (41·3 ppm) copper and 1.0 mm (65·4 ppm) zinc. Egg respiration was also inhibited by copper with 50% inhibition occurring at 0.9 mm (57·2 ppm) whereas zinc inhibited egg respiration by only 30% at 1·5 mM (98·1 ppm).In buffered seawater, both copper and zinc produced an inhibitory effect on sperm respiration. However, with copper the inhibition was much reduced whereas zinc had tthe same inhibitory effect as in unbuffered seawater. In contrast to unbuffered seawater, egg respiration was stimulated by copper but zinc produced a comparable respiratory inhibition. The uptake of both copper and zinc in sperm and egg in buffered seawater increased with increasing metal concentration. Metal ion uptake, when expressed as ng ions μl cell volume^?1, was approximately 3-fold greater in the sperm than in the eggs and in both cases zinc uptake exceeded copper uptake.At a seawater pH of < 7·5, in the absence of copper or zinc, the respiration of both egg and sperm was inhibited. The maximum levels of inhibition at pH 5·4 were 15% and 30% for sperm and egg, respectively.The differing actions of copper on egg respiration in buffered and unbuffered seawater appear to be due to enhanced copper uptake in unbuffered seawater.     

The Energetic and Nutritional Contribution of Glucose and Glycine Taken up from Natural Sea Water by Adult Marine Mussels

Abstract. The removal of glycine and glucose by freshly collected adult Mytilus edulis was determined using radiolabeled substances added to ambient concentrations of dissolved organic materials in freshly collected natural sea water. Uptake rates were calculated for substrate concentrations of 0.5 μM glycine and 1.0 μM glucose and were compared with the animals' energy and nitrogen demands as measured by oxygen consumption and ammonia excretion rates. Respiration and ammonia excretion rates as well as cither glycine or glucose removal were all determined for the same animals. The mean respiration rate was 670μl O₂ g^-1h^-1, the mean ammonia excretion rate 1.95 μg-at NH₄-g^-1 h^-1. The calculated uptake rates were 0.48 μmol -g^-1-h^-1 for free amino acids and 0.44 μmol-g^-1 h^-1 for free simple sugars. Such uptake rates could have contributed roughly 13% of the mussels' energy requirements or 10% of the mussels' nitrogen requirements assuming the following conditions: 0.5 μM concentration of free amino acids, 1.0μM concentration of simple sugars, uptake of total amino acids at a rate based on a weighted removal rate of glycine, alanine, serine, and glutamic acid, and uptake of simple sugars at a rate equal to that of glucose removal. It is apparent that simple organic substances dissolved in sea water may be of some benefit to mussels, especially when the substances occur in concentrations typical of coastal sea water.     

Method for assessing the chronic toxicity of marine and estuarine sediment-associated contaminants using the amphipod Corophium volutator

Acute sediment toxicity tests do not test key life stage events such as moulting and reproduction and therefore do not reveal the longer-term effects of contaminant exposure. A laboratory method is described for determining the chronic toxicity of contaminants associated with whole sediments. The test is conducted using neonates of the estuarine amphipod Corophium volutator at 15 degrees C, salinity 25 psu and a 12 h light:12 h dark photoperiod. The endpoints are survival and growth after 28 days and survival, growth and reproduction of amphipods upon termination of test i.e. reproduction within all control vessels (ca 75 days). The sediment chronic toxicity test was used to investigate the effects of sediments spiked with environmentally relevant preparations of slightly weathered Alaskan North Slope crude oil, including a water-accommodated-fraction (WAF) and a chemically-dispersed (Corexit 9527) WAF. Sediment oil concentrations were quantified using ultra-violet fluorescence. The amphipods exposed to chemically dispersed oil had higher mortality and lower growth rates than control-, Corexit 9527- and WAF-exposed organisms, resulting in reduced reproduction. The described method supplements the standard acute sediment test and would be particularly useful when long-term ecological effects are suspected but acute tests reveal no significant mortality. The sediment chronic test reported herein has shown that sediment that was not evidently toxic during 10-day acute tests could have population-level effects on sediment-dwelling amphipods.     

Behavioural Effects of Chemically Dispersed Oil and Subsequent Recovery in Diploria strigosa (Dana)

Abstract. Survival and behaviour of the hermatypic coral Diploria strigosa was studied during 6–24 h doses with water-accomodated fractions of chemically dispersed crude oil, and for a subsequent recovery period of 1 month. Experiments utilized a flow-through laboratory dosing procedure and incorporated petroleum hydrocarbon measurements in order to simulate a major but short-term oil spill in shallow subtidal benthic reef environments. Chemically dispersed oil treatments consisted of Arabian Light Crude oil with Corexit 9527 or BP1100WD at 1–20 ppm concentrations of oil.
In general, effects observed were sub-lethal, temporary, and associated with the highest concentrations tested. Responses to the presence of dispersed oil at 20ppm for 24 h included mesenterial filament extrusion, extreme tissue contraction, tentacle retraction and localized tissue rupture. The nature and severity of reactions during the dosing phase varied between colonies and treatments, but colonies typically resumed normal behaviour within 2 h to 4 d of the recovery period. It therefore seems unlikely that observed biological effects would impair long-term viability.     

An experimental marine ecosystem response to crude oil and Corexit 9527: Part 2—Biological effects

Three experimental ecosystems were employed to test the effect of Corexit 9527, with and without Prudhoe Bay crude oil, on the ecology of a temperate pelagic ecosystem. The results indicated that Corexit 9527 alone enhanced biological productivity without changing the structure of the ecosystem. The mixture of Corexit and crude oil caused a major change in the ecology of the ecosystem which resulted in large numbers of bacteria and zooflagellates, but a depression of all other zooplankton phyla.     

Editorial

A flow-through system for exposure of seagrass to pollutants is described. Sea water with dissolved atrazine or PCP (pentachlorophenol) was pumped through a sealed 2-litre volume glass exposure vessel that contained either whole plants or leaves of Thalassia testudinum. Effects of the compounds on oxygen evolution and uptake by leaves were measured after exposure for 40 and 88 h. Rate of oxygen evolution was depressed strongly after 40 h by 1 ppm of atrazine or PCP. The rate of oxygen uptake was slightly depressed by atrazine but strongly depressed by PCP. Photosynthesis/respiration ratios were depressed to less than one by 1 ppm of atrazine or PCP and by 0·5 ppm atrazine. EC₅₀ values based upon depression of oxygen evolution after 40 hours exposure were: atrazine, 0·32 ppm PCP, 0·74 ppm. The data suggest that leaves may recover from, or adapt to, the presence of either toxicant.     

营养盐因子对龙须菜和菊花江蓠氮磷吸收速率的影响

本文在实验室条件下,研究了营养盐因子对龙须菜和菊花江蓠N、P吸收速率的 影响,分别对不同氮和磷的浓度、氮磷比及不同化合态氮对这两种藻类氮、磷吸收速 率的影响进行了实验与分析．结果表明:在适宜范围内,两种海藻对氮、磷的吸收速率 随介质中氮、磷浓度的增加而增加;不同氮磷对两种海藻N吸收速率的影响不存在 显著差异,但对磷吸收速率的影响非常显著,最高对磷吸收速率在介质的氮磷比为 10:1时;海藻对NH4-N的吸收速率随着NO3-N／NH4-N比的降低而增加,对PO43-P的 吸收速率与对NH4-N的吸收相似,而对NO3-N的吸收速率与对NH4-N的吸收相反．     

Enzymatic determination of glucose in marine environments: Improvement and note of caution

Microheterotrophic glucose uptake kinetics (V_{max and [K + Sn] value)} and uptake velocities (v) based on natural glucose concentrations (S_n) suggested that glucose concentrations in the Sapelo Island estuary were overestimated by published enzymatic techniques. Glucose oxidase treatment of water samples before glucose measurement yielded results as much as 50% lower than when this step was omitted. Thin layer chromatography shows that glucose must be separated from other carbohydrates (e.g., fructose) which interfere with the glucose assay.     

Summertime trends in pelagic biogeochemistry at the Porcupine Abyssal Plain study site in the northeast Atlantic

Measurements of nitrate and carbon uptake made in July 2006 in the Northeast Atlantic Ocean are evaluated with reference to the photophysiology of the attendant phytoplankton population. Over the 11-day observation period integrated chlorophyll concentrations and carbon fixation rates decreased by 76% and 60%, respectively. Integrated nitrate uptake decreased by 50% from initial to final rates but was generally less variable than carbon fixation and chlorophyll in the intervening period. Satellite derived estimates of surface chlorophyll concentrations reveal the uptake observations to be coincident with, and subsequent to, a peak in summer time production. Large reductions in diatom and dinoflagellate abundance were also seen at this time, with indications that increased grazing, due to an increase in ciliate abundance, was an important mechanism terminating summertime production in the NE Atlantic. Meanwhile, the presence of consistently low values of Fv/Fm (<0.3), particularly in surface waters, suggests that production occurs, or is inhibited, with suboptimal photochemical efficiency widespread amongst the phytoplankton population. Furthermore, the low values of Fv/Fm were not alleviated by day-to-day variability in macronutrient concentration. The timing of our observations places them within the seasonal period recognised for the widespread phenomena of carbon overconsumption, and we estimate C:N uptake ratios at this time could be as high as 13:1.     

Trace metal uptake by the Chinese mitten crab Eriocheir sinensis: the role of osmoregulation

Changes in salinity affect the bioavailability and consequent uptake of trace metals by euryhaline invertebrates. In many cases, salinity-related effects on metal uptake can be explained by changes in chemical speciation but salinity may also influence uptake indirectly through its action on osmoregulatory mechanisms. Specifically, it can be hypothesised that trace metal uptake may be reduced at salinities approaching the isosmotic point of a species because, at this point, there is reduced activity of ionic exchange pumps. The present study tested this hypothesis using the Chinese mitten crab, Eriocheir sinensis, a hyper-hypoosmoregulator with an isosmotic point around 33 per thousand. Crabs were exposed to radio-labelled cadmium and zinc at 23, 33 and 43 per thousand for 4 days. To eradicate speciation effects, crabs were exposed to the same concentration of the radio-labelled free metal ion (estimated using MineQL computer software) at each salinity. Haemolymph samples were taken daily and radio-labelled metal concentrations were estimated from radioactivity counts and used to provide relative measures of metal uptake. Neither cadmium nor zinc uptake was lowest at the isosmotic point. The uptake of cadmium increased significantly with increase in salinity, while the uptake of zinc showed no significant change with increased salinity. Thus changes in trace metal uptake rates in E. sinensis do not appear to be controlled only by changes in free metal ion concentrations. The different effects of salinity change on the uptake of cadmium and zinc (in the absence of free metal ion change) also indicate that physiological responses to osmotic change alone do not control metal uptake rates for this species.     

Influence of combined changes in salinity and copper modulation on growth and copper uptake in the tropical green macroalga Ulva reticulata

The influence of salinity on growth and Cu uptake in the green macroalga Ulva reticulata collected from the intertidal area in the Western Indian Ocean was studied under controlled laboratory conditions. Exposure concentrations ranged from 5 to 500 μg Cu l⁻¹ at five salinities (ranging 20–40). The accumulation of Cu increased with decreasing salinity, so that the uptake at 500 μg Cu l⁻¹ was approximately 2.7, 2.4 and 2.0 times higher at salinities of 20, 25, and 30 respectively, than uptake at salinity of 35, and with uptake being lowest at salinity of 40. Ulva maintained a positive growth rate over the whole salinity range (20–40), with highest rates at salinity of 35. When exposing to Cu at low salinities (20 and 25), the growth rate of Ulva was strongly inhibited suggesting an increase in toxicity of Cu with decreasing salinity. EC50 and NOEC increased with increase in salinity, implying a reduced Cu toxicity at high salinities. It was concluded that salinity needs to be considered when using macroalgae, such as U. reticulata, as a bioindicator of heavy metals in areas with heavy rainfall, underground fresh water intrusion or in estuaries, as they might accumulate more metals and be more negatively affected.     

微量元素锰对海带生长的效应

近年来,微量元素对植物生长发育的影响日益受到人们的重视。实验证明这类元素不仅对一些农作物如水稻和小麦[1,3,4]等的早期生长有促进作用,而且对一些藻类植物如小球藻[1]等也有同样作用。正是在这种情形下,利用锰离子提高作物生产的实验正在开展中。从已有报导来看,锰肥的施用对不少种植物有明显效应。如当锰量适当时,玉米籽粒的重量可以大为增加[4]。反之如果缺锰,小球藻的氮代谢——尤其是硝酸盐的同化作用过程就会受到影响。这时如果在培养液中增加锰浓度,可见在短时间内,光合作率就能成倍增加。     

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.     

Modeling the biodegradation of phenolic compounds by microalgae

Phenols represent a group of organic pollutants frequently found in many near-shore marine systems. The microbial degradation of phenols, mainly by bacteria and fungi, has been extensively studied both experimentally and theoretically, but only relatively recently the capabilities of some algae for phenols biodegradation gained interest. The biodegradation of phenols by microalgae occurs only under aerobic conditions. In this paper, a dynamic energy budget model is proposed for describing aerobic biodegradation of phenolic compounds by microalgae and qualitatively validated against experimental data. A microalgal cell has the ability to produce biomass via the autotrophic assimilation (uptake of light and dissolved inorganic carbon), the heterotrophic assimilation (uptake of dissolved organic carbon) and, to a lesser extend, via the biodegradation of phenols. The rules of synthesizing units are used for the uptake and interactions of substrates and for the merging of assimilates. The model is capable of making predictions under oxygen and carbon (inorganic and organic) limiting conditions. Model predictions cover a wide range of experimental evidence, but also give a possible explanation for the inhibition of bioremoval of phenols in the presence of glucose. The dissolved oxygen profiles numerically observed show low oxygen concentration during the intermediate phase of the biodegradation process and a rapid increase after the consumption of the phenolic compound, indicating that lack of oxygen could be a limiting factor for the biodegradation of phenols. The presence of glucose increases the specific growth rate but decreases the specific biodegradation rate of the phenolic compound. Model analysis suggests that this inhibition may be due to the competition for oxygen between glucose and phenol assimilation. In general, the balance between the benefits and costs of the different types of assimilation determines the microalgal growth rates as well as the phenol biodegradation rates. Being based on general assumptions, the model can be applied to the biodegradation of a wide variety of aromatic compounds.     

Relative importance of hydrocarbon pollutants, salinity and tidal height in colonization of oyster reefs

The relative effects of hydrocarbon pollutants, salinity and tidal height on the invertebrates and fish that inhabit oyster reefs were studied along the Louisiana Gulf of Mexico coastline. Dried oyster shell (cultch) was first exposed to crude oil in the laboratory. In a series of experiments, plastic trays filled with control and oil-exposed cultch were then deployed at two locations differing in salinity, in two seasons and at two tidal levels. In experiments on hydrocarbon, salinity, and seasonal effects, trays were colonized for one month. To examine the effects of colonization time, half of the trays were retrieved after two and the rest after five weeks. Salinity dramatically affected oyster reef assemblages, with species richness and total abundance halved at the estuarine site. Hydrocarbon effects were less prominent, whether cultch was dosed with light or heavy crude oil. The sub-tidal site had higher colonization rates, but colonization interval did not affect colonization, and seasonal differences occurred only at the higher-diversity, sub-tidal site. To determine effects of cleaners, Corexit 9580 was applied alone and with oil on cultch, and trays were colonized for one month. At high concentrations, the cleaner ameliorated hydrocarbon effects. In general, hydrocarbon effects were less prominent than salinity and aerial exposure in explaining colonization of oyster reef assemblages. Gas chromatography/mass spectrometry analysis of oyster shells after one month immersion revealed considerable losses of oil, especially with higher flow at the inter-tidal site. Sediment on shell also diluted oil. We argue that oyster reef assemblages should recover from small-scale spills, unless they occur during periods of reproduction and dispersal.     

Benthic mineralization rates at two locations in the southern North Sea

Benthic oxygen uptake, sulphate reduction and benthic bacterial production were measured at two contrasting locations in the southern North Sea: the shallow and turbulent Broad Fourteens area in the Southern Bight, and the deeper Oyster Grounds, a deposition area, where thermohaline stratification occurs during summer. Oxygen uptake and sulphate reduction showed a clear seasonal pattern in the Broad Fourteens area, indicating a supply of carbon to the benthic system that is closely related to the standing stock of carbon in the water column. This close benthic-pelagic coupling is probably due to the influence of the tide in this part of the North Sea, which keeps the water column permanently mixed. At the Oyster Grounds, no seasonal pattern was observed. Peaks in oxygen uptake and sulphate reduction were found in winter. Irregularly occurring events, such as storms and fishery-related activities, are likely to affect the benthic mineralization patterns in this area. Annual benthic carbon mineralization rates estimated from oxygen uptake rates were 44 gC·m⁻² at the Broad Fourteens, and 131 gC·m⁻² at the Oyster Grounds, of which 26 and 28%, respectively, could be attributed to sulphate reduction (assuming an annual sulphide reoxidation rate of 100%). Although sulphate reduction rates in the southern North Sea are higher than previously suggested, aerobic respiration is the most important pathway for benthic carbon mineralization at the stations visited. Production rates of benthic bacterial carbon measured with labelled leucine were much higher than carbon mineralization rates based on oxygen uptake or sulphate reduction. This may either imply a very high bacterial carbon conversion efficiency, or point to shortcomings in the accuracy of the techniques. A critical evaluation of the techniques is recommended.     

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.     

Effect of environmental factors on photodegradation of polycyclic aromatic hydrocarbons (PAHs) in the water-soluble fraction of Kuwait crude oil in seawater

Photodegradation of PAHs in the water-soluble fraction of Kuwait crude oil in seawater was investigated under various environmental factors (temperature, light intensity, oxygen levels and presence of a sensitizer) in laboratory conditions. All factors investigated had significant effect on the degradation rates of PAHs. At 15 °C almost all PAHs optimally degraded at an oxygen level of 4 ppm. For lower molecular weight PAHs a light intensity of 500 W/m² in the presence of the sensitizer worked well. Higher molecular weight PAHs degraded at faster rates at a light intensity 750 W/m². At 30 °C, most of the PAHs degraded optimally at an oxygen level of 0 ppm and light intensity of 500 or 750 W/m² in presence of the sensitizer. At 40 °C, most of PAHs degraded optimally at low oxygen concentrations (0 and 4 ppm) and a light intensity of 500 W/m² in the presence of the sensitizer. Linear regression indicated that for most of the compounds, light intensity had the greatest effect on degradation rates.     

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.