Development of rapid ammonium uptake during starvation of batch and chemostat cultures of the marine diatom Thalassiosira pseudonana

Cell nitrogen quotas and uptake rates following ammonium additions were measured during ammonium-limited growth transients obtained by starving batch and chemostat cultures of Thalassiosira pseudonana (Clone 3 H). During starvation, cell quotas decreased by more than 50% in batch cultures. In chemostat cultures, the drop in cell quota during starvation decreased with dilution rate, from more than 50% at 1.45 d^-1, to less than 10% at 0.22 d^-1. Minimal levels of 3 to 4×10^-2 pg-at. N cell^-1 were reached after 24 h starvation in both batch and chemostat cultures. Uptake rates over the first minute of perturbation experiments were 3 times the long-term (10 to 30 min) rates. In batch cultures, specific uptake rates increased from 4 d^-1 to 20 d^-1 after 24 h starvation. Uptake rates per cell were independent of starvation time and dilution rate in chemostat cultures, but lower in non-starved batch cultures. The implications of these data for models of phytoplankton growth are discussed: the data support models which predict a depression in average growth rates when diatoms encounter microscale patches in oligotrophic environments.     

Effect of nitrogen supply on nitrogen uptake,accumulation and assimilation in Porphyra perforata (Rhodophyta)

Porphyra perforata J. Ag. was collected from a rocky land-fill site near Kitsilano Beach, Vancouver, British Columbia, Canada and was grown for 4 d in media with one of the following forms of inorganic nitrogen: NO ₃ ^- , NH ₄ ⁺ and NO ₃ ^- plus NH ₄ ⁺ and for 10 d in nitrogen-free media. Internal nitrogen accumulation (nitrate, ammonium, amino acids and soluble protein), nitrate and ammonium uptake rates, and nitrate reductase activity were measured daily. Short initial periods (10 to 20 min) of rapid ammonium uptake were common in nitrogen-deficient plants. In the case of nitrate uptake, initial uptake rates were low, increasing after 10 to 20 min. Ammonium inhibited nitrate uptake for only the first 10 to 20 min and then nitrate uptake rates were independent of ammonium concentration. Nitrogen starvation for 8 d overcame this initial suppression of nitrate uptake by ammonium. Nitrogen starvation also resulted in a decrease in soluble internal nitrate content and a transient increase in nitrate reductase activity. Little or no decrease was observed in internal ammonium, total amino acids and soluble protein. The cultures grown on nitrate only, maintained high ammonium uptake rates also. The rate of nitrate reduction may have limited the supply of nitrogen available for further assimilation. Internal nitrate concentrations were inversely correlated with nitrate uptake rates. Except for ammonium-grown cultures, internal total amino acids and soluble protein showed no correlation with uptake rates. Both internal pool concentrations and enzyme activities are required to interpret changes in uptake rate during growth.     

Interactions between nitrate and ammonium uptake: variation with growth rate,nitrogen source and species

The interaction between nitrate and ammonium uptake was examined as a function of preconditioning growth rate and nitrogen source by adding nitrate, ammonium, or both to nitrogen-sufficient,-deficient, and-starvedSkeletonema costatum (Grev.) Cleve and nitrogen-deficientChaetoceros debilis Cleve. By simultaneously measuring the internal accumulation of intermediates of nitrogen assimilation and the rates of nitrogen assimilation, the metabolic control of nitrogen uptake could be assessed. After the simultaneous addition of nitrate and ammonium to culture, both nitrate and ammonium uptake rates were decreased in comparison with the rates observed when each was added alone, although nitrate uptake was usually decreased more than ammonium uptake. Since both nitrate and ammonium uptake rates vary with time, preconditioning growth conditions, nitrogen sources present, and species, it was necessary to use several different indices to quantify inhibition. In general, ammonium inhibition of nitrate uptake inS. costatum was greatest in cultures preconditioned to ammonium and those at low growth rates, whereas ammonium uptake was inhibited most in cultures preconditioned to nitrate. In nitrogen-deficientC. debilis, nitrate uptake was more inhibited by ammonium, but uptake returned to normal rates more quickly than inS. costatum, whereas inhibition of ammonium uptake was similar. These results explain why the interaction between nitrate and ammonium uptake in the field can be so variable. Inhibition of uptake is not controlled by internal ammonium or total amino acids, nor is it related to the inability to reduce nitrate. Instead, inhibition must be determined in part by the external concentration of nitrogen compounds and in part by some intermediate(s) of nitrogen assimilation present inside the cell.Bigelow Laboratory Contribution No. 82022     

Response of marine phytoplankton to nitrogen deficiency: Decreased nitrate uptake vs enhanced ammonium uptake

The uptake of nitrate and ammonium was measured separately in uni-algal, nitrogen-deficient cultures of four species of marine phytoplankton. Nitrogen-deficient phytoplankton took up ammonium at initial rates which greatly exceeded those measured for nitrogen-sufficient phytoplankton. However, nitrate uptake by nitrogendeficient cultures was generally much slower than either nitrate or ammonium uptake by nitrogen-sufficient cultures or ammonium uptake by nitrogen-deficient cultures. Considerable species differences were observed in the degree to which nitrogen deficiency increased ammonium uptake or decreased nitrate uptake. Loss of ability to take up nitrate, but enhanced ability to take up ammonium, as a result of nitrogen deficiency may be an adaptation to the different mechanisms by which nitrate and ammonium are supplied to the euphotic zone. In areas with an intermittent supply of nitrogen, changes in the ability of some species to take up nitrogen as a result of nitrogen starvation will influence species composition and complicate interpretations of measurements of nitrogen uptake.Contribution no. 1249 from the Department of Oceanography, University of Washington, and contribution no. 82006 from the Bigelow Laboratory for Ocean Sciences     

Nitrate metabolism in sediments from seagrass (Zostera capricorni) beds of Moreton Bay,Australia

The fate of nitrate in sediments from seagrass (Zostera capricorni Aschers.) beds of Moreton Bay on the subtropical eastern coast of Queensland, Australia, was investigated. Added nitrate was metabolised at rates of 0.4 to 3.4 g N cm^-3 d^-1 when sediments were incubated under anaerobic conditions with a large excess of nitrate. The potential rate of nitrate utilization was as rapid in sediments from subtidal bare areas as from adjacent seagrass beds. Ammonium was produced rapidly from¹⁵N-nitrate by microbial action in all the subtidal sediments examined. After 12 h of incubation, 13 to 28% of the¹⁵N initially added as labelled nitrate was detected as labelled ammonium in the sediments. Denitrification, although not measured directly, appeared to be a relatively minor fate of nitrate. Benthic microbes took up large amounts of¹⁵N but only after a delay of 6 h; this pattern could have been due to induction and synthesis of the enzymes necessary for nitrate uptake, and the assimilation of labelled ammonium. Under field conditions, assimilation by seagrasses and denitrification by bacteria were probably not significant sinks for nitrate in comparison with uptake by benthic microbes and dissimilatory reduction to ammonium.     

Nitrate uptake by the reef coral Diploria strigosa: effects of concentration, water flow, and irradiance

The effects of several environmental variables on net nitrate uptake by the scleractinian coral Diploria strigosa were investigated under controlled flow conditions. D. strigosa exhibited nitrate uptake rates ranging from 1 to 5 nmol cm⁻² h⁻¹ at ambient concentrations of 0.1–0.3 μM that are typical of oligotrophic reefs such as Bermuda. Net uptake ceased at approximately 0.045 μM. The uptake was positively correlated with concentration up to a saturation concentration of approximately 3 μM. The uptake was also positively correlated with water velocity at 1 μM, but not at 6 μM, suggesting diffusional limitation at low concentrations and kinetic limitation at higher concentrations. Nitrate uptake by D. strigosa was not affected by light intensity or time of day, but was almost completely inhibited by 48 h exposure to ammonium levels found on many reefs.     

Synergistic effects of nitrate and ammonium ion on the growth and uptake kinetics of Monochrysis lutheri in continuous culture

Continuous-culture results for Monochrysis lutheri grown on 12 h light-12 h dark cycles with a spectrum of ratios of nitrate and ammonium serving as limiting nutrients are compared with continuous light, exclusively nitrate, and exclusively ammonium-limited data for this species. The diel effects of the light regime on the maximum specific uptake rate are examined for both nitrate and ammonium. Synergistic effects on uptake by various initial concentrations of these two nutrients are presented. Preconditioning with light-dark cycles did not affect maximum uptake rate, but preconditioning on a combination of nitrate and ammonium gave much lower uptake rates than those observed for populations preconditioned on either nutrient exclusively. The implications of high maximum specific-uptake rates compared to maximum specific-growth rates in terms of the range of nitrate and ammonium ion concentrations associated with nutrient limitation are reviewed.Hawaii Institute of Marine Biology Contribution No. 477.     

Marine diatoms grown in chemostats under silicate or ammonium limitation. I. Cellular chemical composition and steady-state growth kinetics of Skeletonema costatum

Skeletonema costatum was grown in chemostats under ammonium or silicate limitation to examine its growth kinetics and changes in cellular chemical composition at different steady-state growth rates. When the relationship between the effluent limiting substrate concentration and steady-state growth rates was examined, deviations from the simple hyperbolic form of the Monod growth equation were noted at low and high dilution rates. The data from the plot of growth rate and substrate concentration were divided into 4 regions and the relationship of these region to cell quota is discussed. Two physiological states were identified. All populations grown at D<0.05 h^-1, regardless of the size of the cells or the magnitude of Q, exhibited a maximal growth rate of approximately 0.05 h^-1, while populations grown at higher dilution rates (D>0.06 h^-1 to 0.14 h^-1). The maximal value of growth rate is obtained only in cultures grown at very high dilution rates where nutrient shift-up appears to occur, the cell quota approaches a maximum and the heterogeneous cell population becomes more homogeneous.Contribution No. 881 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. This paper represents a portion of two dissertations submitted by P.J.H. and H.L.C. to the Department of Oceanography, University of Washington, Seattle, in partial fulfillment of the requirements for the Ph.D. degree.     

Nitrogen uptake kinetics in three year-classes of Laminaria groenlandica (Laminariales: Phaeophyta)

Nitrate and ammonium uptake rates were measured for three year-classes of the perennial macrophyte Laminaria groenlandica Rosenvinge, collected from nitrogen-depleted waters in Barkley Sound, British Columbia, Canada, in summer 1981. A time course of uptake rate revealed that ammonium uptake was high during the first hour and then decreased for all three year-classes; the opposite pattern was exhibited for the time course of nitrate uptake rate. Nitrate uptake rate increased linearly with nitrate concentration up to the highest level tested (60 M). The nitrate uptake rate of first-year plants was three times higher than second- and third-year plants; ammonium uptake rates showed similar patterns to those for nitrate. The interaction between nitrate and ammonium was examined for first-year plants. Nitrate and ammonium were taken up simultaneously and uptake rates were identical and equal to uptake rates when only nitrate or ammonium was present in the medium. Therefore, first-year plants are able to take up twice as much inorganic nitrogen per unit time when both nitrate and ammonium are present. First-year plants showed significant diel periodicity in ammonium uptake rates, whereas second- and third-year plants showed no periodicity in nitrate or ammonium uptake rates.     

Short-term interaction between nitrate and ammonium uptake inThalassiosira pseudonana: Effect of preconditioning nitrogen source and growth rate

The effect of preconditioning nitrogen source and growth rate on the interaction between nitrate and ammonium uptake was determined inThalassiosira pseudonana (Clone 3H). A new method, using cells on a filter (Parslow et al. 1985), allowed continuous measurement of uptake from 0.5 to 9 min after the addition of nitrate, ammonium, or both, with no variation in concentration during the course of the experiment. For each preconditioning N source and growth rate, a series of uptake experiments was conducted, including controls with only nitrate or only ammonium, and others with different combinations of concentrations of nitrate and ammonium. For the first time, preference for ammonium was separated from inhibition of nitrate uptake by ammonium. Ammonium was the preferred N source, i.e. if nitrate and ammonium were presented separately, ammonium uptake rates exceeded nitrate uptake rates. Preference for ammonium varied with both preconditioning N source and growth rate. Inhibition of nitrate uptake by ammonium, determined by comparing nitrate uptake in the presence and absence of ammonium, was observed at ammonium concentrations > 1µM, but was rarely complete. Inhibition of nitrate uptake by ammonium was less in the ammonium-limited culture than in the cultures growing on nitrate, but invariant with growth rate in the nitrate-grown cultures. Below 1µM ammonium, nitrate uptake was often stimulated and rates exceeded those in the controls without ammonium. Ammonium uptake was not inhibited by the presence of nitrate.T. pseudonana fits the classical view of the interaction between nitrate and ammonium uptake in some respects, such as preference for ammonium, and inhibition of nitrate uptake by ammonium concentrations > 1µM. However, at ammonium concentrations typical of most marine environments, nitrate uptake occurs at rapid rates. In other respects, N uptake inT. pseudonana deviates from the classical view in the following ways: (1) stimulation of nitrate uptake by low concentrations of ammonium; (2) lack of inhibition of nitrate uptake by ammonium at low nitrate concentrations; and (3) variation in preference and inhibition with preconditioning, which is markedly different for other species. Because of the apparent enormous species variation in the interaction between nitrate and ammonium uptake and the lack of detailed information for a variety of species, it is difficult to generalize about the effect of ammonium on nitrate uptake, especially in the field, where prior N availability and species composition are not usually addressed.     

Partitioning of nitrogen and carbon in cultures of the marine diatom Thalassiosira fluviatilis supplied with nitrate,ammonium, or urea

Small or negligible differences in growth rates, average cell size, yields in cell numbers and total cell volumes were found in cultures of Thalassiosira fluviatilis inriched with nitrate, ammonium, or urea. Intracellular pools of unassimilated nitrate, nitrate, and ammonium were found in nutrient-rich conditions, but urea was not accumlated internally. Nitrogen assimilation into organic combination rather than nitrogen nutrient uptake was a critical rate-limiting step in nitrogen utilization. The free amino acid pool, protein, lipid-associated nitrogen, pigments, and total cell nitrogen were all highest in young or mature phase cells and decreased with age in senescent cells, whereas chitan, lipid, carbohydrate, and total cellular carbon all continued to increase during senescence. Dissolved organic nitrogen compounds accumulated in the medium only during senescence. C:N and lipid:protein were sensitive indicators of nitrogen depletion and age in T. fluviatilis.     

The rapid response of the marine diatom Skeletonema costatum to changes in external and internal nutrient concentration

A nitrogen-deficient batch culture of the marine diatom Skeletonema costatum, when resupplied with a mixture of nitrate and ammonium, showed an initial enhanced nitrate uptake rate leading to a large internal concentration (pool) of nitrate. Following this initial nitrate uptake event, nitrate uptake ceased, and nitrate assimilation was inhibited until the ammonium present was used. At this point, nitrate uptake resumed and nitrate assimilation began. No internal ammonium pool was observed during nitrate utilization, but a large nitrate pool remained throughout the utilization of external nitrate. The internal nitrate pool decreased rapidly after exhaustion of nitrate from the culture medium, but growth of cellular particulate nitrogen continued for about 24 h. A mathematical simulation model was developed from these data. The model cell consisted of a nitrate pool, ammonium pool, dissolved organic nitrogen pool, and particulate nitrogen. It was found that simple Michaelis-Menten functions for uptake and assimilation gave inadequate fit to the data. Michaelis-Menten functions were modified by inclusion of inhibitory and stimulatory feedback from the internal pools to more accurately represent the observed nutrient utilization.     

Sinking-rate response of natural assemblages of temperate and subtropical phytoplankton to nutrient depletion

Phytoplankton assemblages were collected during spring blooms in 1982 in Washington State and in Hawaii. Sinking rate responses of these assemblages were examined under nitrate, phosphate, and silicate depletion. Ambient nutrient concentrations, chlorophyll concentrations, photosynthetic rates, sinking rates, and floristic compositions were determined. Under nutrient-replete conditions, the temperate assemblage, composed primarily of large centric diatoms, had a sinking rate of 0.96 m d^-1; sinking rates did not change appreciably over 4 d without nitrate. Without phosphate or silicate, the sinking rates remained constant for 3 d and then increased after biomass indices began to decline. These findings illustrate the potential importance of phosphate or silicate depletion to the sedimentation of spring-bloom diatom populations. The subtropical assemblage, composed primarily of diatoms, coccolithophorids, and dinoflagellates, had an initial sinking rate of 0.22 m d^-1 and did not display substantial sinking rate changes in the absence of nitrate, phosphate or silicate. Floristic data consistently showed a proliferation of pennate diatoms, which had lower settling rates than centric diatoms. Growth and sedimentation patterns indicated a competitive advantage for pennate diatom components of subtropical assemblages; this in turn may limit phytoplankton sedimentation losses in such ecosystems.     

Phosphorus metabolism of oceanic dinoflagellates: phosphate uptake,chemical composition and growth of Pyrocystis noctiluca

The phosphorus metabolism of Pyrocystis noctiluca Murray (Schuett) 1886 has characteristics which may enhance its potential for success in orthophosphate impoverished waters. The steady-state phosphate uptake rates were equal in the light and dark, and were directly proportional to both the phosphorus cell quota and the cell division rate. In contrast, nutrient-saturated uptake rates were multiphasic, faster in the light than the dark, 2 to 4 orders of magnitude greater than steady-state rates, and were inversely proportional to both the phosphorus cell quota and the cell division rate. These uptake characteristics suggest that P. noctiluca may take up phosphate coincidently at their typically low ambient concentrations as well as to exploit episodic nutrient events in nature. Cell division rates were a hyperbolic function of the ambient orthophosphate concentration. The shortest doubling time was 8.7 d, the phosphate concentration at half the maximum division rate was 0.15 M and the threshold, concentration for cell division was ca 0.05 M PO ₄ ^3- . Division rates of P. noctiluca in the ocean are much faster than predicted from the measured ambient orthophosphate concentrations. Since this dinoflagellate has high naturally occurring alkaline phosphatase activities, and can utilize organic-P compounds, we suggest that organic-P can be as important as orthophosphate in supporting the observed division rates of P. noctiluca in the sea.     

Effects of host feeding and dissolved ammonium on cell division and nitrogen status of zooxanthellae in the hydroid Myrionema amboinense

There is a relationship between host feeding, nitrogen status and mitotic activity of zooxanthellae symbiotic with the marine hydroid Myrionema amboinense. Decreases in the mitotic index of zooxanthellae in starved M. amboinense, and in internal pool sizes of glutamine and glutamate, amino acids involved in ammonium assimilation via the glutamine synthetase-glutamate synthase (GS/GOGAT) pathway, were partially restored by addition of ammonium chloride to seawater in which hydroids were incubated. Levels of glutamine were more sensitive to host starvation than levels of glutamate, resulting in a decrease in the glutamine: glutamate molar ratio to that found in zooxanthellae cultured on nitrate. Hydroids starved for 5 d and then incubated in different concentrations of ammonium chloride showed a positive correlation between ammonium concentration and mitotic index of their symbiotic zooxanthellae. Host starvation caused a decrease in perturbation of levels of glutamine and glutamate during ammonium assimilation, as well as decreases in rates of assimilation of [¹⁴C]-leucine into TCA-insoluble protein, and in photosynthetic incorporation of [¹⁴C]-bicarbonate. These observations suggest that host starvation reduces nitrogen supply to the zooxanthellae, causing nitrogen stress to the symbionts and reduction in metabolic processes associated with nitrogen assimilation and photosynthesis as well as with cell division.     

Marine diatoms grown in chemostats under silicate or ammonium limitation. III. Cellular chemical composition and morphology of Chaetoceros debilis,Skeletonema costatum,and Thalassiosira gravida

Three marine diatoms, Skeletonema costatum, Chaetoceros debilis, and Thalassiosira gravida were grown under no limitation and ammonium or silicate limitation or starvation. Changes in cell morphology were documented with photomicrographs of ammonium and silicate-limited and non-limited cells, and correlated with observed changes in chemical composition. Cultures grown under silicate starvation or limitation showed an increase in particulate carbon, nitrogen and phosporus and chlorophyll a per unit cell volume compared to non-limited cells; particulate silica per cell volume decreased. Si-starved cells were different from Si-limited cells in that the former contained more particulate carbon and silica per cell volume. The most sensitive indicator of silicate limitation or starvation was the ratio C:Si, being 3 to 5 times higher than the values for non-limited cells. The ratios Si:chlorophyll a and S:P were lower and N:Si was higher than non-limited cells by a factor of 2 to 3. The other ratios, C:N, C:P, C:chlorophyll a, N:chlorophyll a, P:chlorophyll a and N:P were considered not to be sensitive indicators of silicate limitation or starvation. Chlorophyll a, and particulate nitrogen per unit cell volume decreased under ammonium limitation and starvation. NH₄-starved cells contained more chlorophyll a, carbon, nitrogen, silica, and phosphorus per cell volume than NH₄-limited cells. N:Si was the most sensitive ratio to ammonium limitation or starvation, being 2 to 3 times lower than non-limited cells. Si:chlorophyll a, P:chlorophyll a and N:P were less sensitive, while the ratios C:N, C:chlorophyll a, N:chlorophyll a, C:Si, C:P and Si:P were the least sensitive. Limited cells had less of the limiting nutrient per unit cell volume than starved cells and more of the non-limiting nutrients (i.e., silica and phosphorus for NH₄-limited cells). This suggests that nutrient-limited cells rather than nutrient-starved cells should be used along with non-limited cells to measure the full range of potential change in cellular chemical composition for one species under nutrient limitation.Contribution No. 943 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Nutrient starvation effects on the allelochemical potency of Alexandrium tamarense (Dinophyceae)

Batch culture experiments were performed to investigate potential effects of nutrient starvation on the allelochemical potency of the toxic dinoflagellate Alexandrium tamarense. Triplicate cultures with reduced nitrate (−N) or phosphate (−P) seed were compared to nutrient-replete (+N+P) cultures. Total depletion of the dissolved inorganic limiting nutrient, reduced cell quotas, changed mass ratios of C/N/P and reduced cell yield clearly indicate that treatment cultures at stationary phase were starved by either N or P, whereas growth cessation of +N+P cultures was probably due to carbon limitation and/or a direct effect of high pH. Pulsed addition of the limiting nutrient allowed −N and −P cultures to resume growth. Lytic activity of A. tamarense as quantified by a Rhodomonas bioassay was generally high (EC₅₀ around 100 cells mL⁻¹) and was only slightly modulated by growth phase and/or nutrient starvation. Lytic activity per cell increased with time in both +N+P and −P cultures but not −N cultures. P-starved stationary-phase cells were slightly more lytic than +N+P cultures, but this difference may be due to increased cell size and/or accumulation of extracellular compounds. In conclusion, only slight changes but no general and major increase in lytic activity in response to nutrient starvation was observed.     

Nitrogenous nutrition in the euphotic zone of the Central North Pacific Gyre

The uptake rates of the three nitrogen compounds ammonium, nitrate, and urea were measured in the oligotrophic North Central Pacific Gyre in August–September 1985. The measurements were performed by using ¹⁵N-labelled substrates and incubating for short-time periods (3 to 4 h) under simulated in situ conditions. Ambient concentrations of the nitrogenous nutrients were generally below 0.10 mol l^-1. The average total daily nitrogen uptake rate, integrated over the euphotic zone, was 12.5 mmol N m^-2 d^-1. Diel studies in the upper water mass resulted in a calculated phytoplankton growth rate of 1.3 d^-1. Ammonium was the dominating nutrient, accounting for on the average 54% of the total nitrogen uptake, while urea uptake represented 32% and nitrate 14%. Ammonium uptake rates at a coastal station off the Hawaiian Islands were very close to the rates found at the oceanic station. Organisms <3 m dominated the nitrogen assimilation, being responsible for about 75% of the ammonium uptake. The nitrogen uptake rates in this study seem to be higher than those found by earlier investigations in the area, but correlated well with other productivity measurements performed during the same cruise.     

Seasonal variations in the utilization of ammonium and nitrate by photoplankton in Vineyard Sound,Massachusetts, USA

The nitrogenous nutrition of the phytoplankton in Vineyeard Sound, Massachusetts, USA was investigated over a 15-month period. Highest rates of ammonium uptake were observed immediately prior to, or during, the diatom bloom periods, and with one exception were found in the <10 m size class. The saturating rate of ammonium uptake correlated well with temperature and gave Q₁₀ values of 2.6–3.2; correlations with ambient solar irradiation were not nearly so clear. Uptake rates of ammonium exceeded those of nitrate except during the winter bloom of the diatom Rhizosolenia delicatula; yet calculation of the f ratio revealed that nitrate was relatively important in the nitrogenous nutrition of the phytoplankton throughout the year.Contribution no. 5096 from the Woods Hole Oceanographic Institution     

Adaptations of Gracilaria pacifica (Rhodophyta) to nitrogen procurement at different intertidal locations

Nitrogen uptake, assimilation and accumulation were studied in three populations of Gracilaria pacifica Abbott in Bamfield Inlet, British Columbia, Canada, over three summers, 1979–1981. Two of these populations were in the intertidal one high and one low, and the third was a subtidal cultured population. Nitrate uptake rates, internal nitrate content and nitrate reductase activities were highest in the low intertidal population. Time-courses of uptake and uptake kinetics were studied. Both nitrate and ammonium were taken up simultaneously. Thalli from the high-intertidal population showed enhanced nitrate and ammonium uptake following mild desiccation, and greater tolerance to desiccation in terms of maintaining nitrogen uptake after severe desiccation. Transplants were made to determine the effect of intertidal height and geographic location on responses to desiccation, nitrogen uptake, assimilation and accumulation. Nitrate and ammonium uptake rates were dependent on intertidal height and geographic location. Transplanting up the intertidal increased nitrate uptake and nitrate reductase activity, but decreased the nitrate content of the thalli. There were few significant differences in ammonium uptake rates, and ammonium, amino acid, and soluble-protein content of the various populations. All high-intertidal populations, transplanted or natural, showed enhanced nitrate uptake rates following desiccation. Enhanced ammonium uptake rates following desiccation were restricted to the high-intertidal thalli in only one geographic location. Tolerance to higher levels of desiccation also appeared to be intertidal height-dependent, but required more than five weeks to fully develop or disappear.