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.     

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.     

Nitrogen uptake and growth of the germlings and mature thalli of Fucus distichus

Fucus distichus L. was collected near Vancouver, Canada, in late fall and early winter, 1981. The effects of the forms of nitrogen (nitrate, ammonium or urea) and periodic exposure to air on growth, rhizoid development and nitrogen uptake in germlings was investigated. Gamete release, fertilization, germination and germling growth had no requirement for a specific form of nitrogen. Periodic exposure to air increased secondary rhizoid development twofold. Nitrate and ammonium uptake rates of the germlings were higher than for the mature thalli (20 to 40 times for nitrate and 8 times for ammonium), while the halfsaturation constant (K _s) values for nitrate were similar (1 to 5 M). The germlings showed saturable uptake kinetics but the mature thalli did not. When germlings were exposed to air it caused a 70% decrease in nitrate uptake, but not change in ammonium uptake. Ammonium uptake in the mature thalli was proportional to the ambient ammonium concentration. Nitrate uptake in the mature thalli appeared to follow saturation kinetics at low nitrate concentrations, but showed a non-saturable component at concentrations greater than 10 M. Presence of ammonium inhibited nitrate uptake by the mature plants but not by the germlings.     

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.     

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     

The nitrogen supply to intertidal eelgrass (Zostera marina)

The uptake of ammonium and nitrate by eelgrass (Zostera marina L.) was studied in two-compartment chambers. The plants were collected in 1992 from a population growing on a tidal flat in the S.W. Netherlands. They were incubated under conditions which reflected field conditions; this implied the use of natural seawater and sediment porewater as incubation media. In all six experiments, carried out over the course of a major part of the growing period (from July to the end of September), ammonium appeared to be much more important as a source of nitrogen than nitrate. The largest part was taken up by the leaves: uptake of ammonium by the leaves accounted for 68 to 92% of total plant nitrogen uptake. The uptake of nitrogen compounds by the root-rhizome system represented only 4 to 30% of total plant uptake. Thus, at least during flood tide, the leaves play the major role in nitrogen uptake in this intertidal population. During ebb tide, most of the plants are submerged in very shallow tidepools. It is suggested that during this phase of the tidal cycle, influx of porewater ammonium into the tide-pool water may enable the leaves to exploit local sediment resources.     

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.     

Effect of bryozoan colonization on inorganic nitrogen acquisition by the kelps Agarum fimbriatum and Macrocystis integrifolia

The effect of bryozoan colonization on inorganic nitrogen acquisition by Agarum fimbriatum Harv. and Macrocystis integrifolia Bory., collected from the west coast of Vancouver Island, British Columbia, Canada, was examined in laboratory experiments during June and July 1992. Pieces of kelp blades that were completely covered on one side by the bryozoans Lichenopora novae-zelandiae Busk or Membranipora membranacea, L., or uncolonized (clean treatment), were used to estimate the rate at which nitrate and ammonium were removed from the surrounding seawater. In addition, the rate of ammonium excretion by bryozoans isolated from their associated kelp was measured and also estimated from the results of the uptake experiments. Values obtained were used to estimate the contribution of ammonium excreted by bryozoans to the total amount of inorganic nitrogen available to the associated kelp. Both bryozoan species reduced the ability of the associated kelp to remove nitrate and ammonium from seawater but provided a source of ammonium to the kelp through excretion. The nitrogen status of colonized and clean kelp disks was determined from the ratio of total particulate carbon to total particulate nitrogen (C:N ratio). The C:N ratios for A. fimbriatum colonized with either L. novae-zelandiae or M. membranacea were similar (C:N=12 to 14), and differences between colonized and clean treatments were not significant. For A. fimbriatum, therefore, the C:N ratio indicates that this species was not nitrogen limited at the time of the present study. In contrast, both colonized and clean disks of M. integrifolia were nitrogen limited, but colonized disks (C:N=19) were significantly less limited by nitrogen than clean disks (C:N=29). Results are discussed in relation to the different environments inhabited by both kelp species and are consistent with the hypothesis that ammonium excreted by bryozoans was an important source of inorganic nitrogen to M. integrifolia, but not to A. fimbriatum, at the time of the study.     

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     

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.     

Regulation of nitrate and ammonium uptake in the Greenland Sea

The uptake of nitrate and ammonium was investigated experimentally during early spring 1989 in the Greenland Sea, with particular attention placed on the roles of irradiance, nitrogen concentrations and nitrateammonium interactions. The phytoplankton assemblage was dominated by the colonial prymnesiophyte Phaeocystis pouchetii. Nitrate concentrations ranged from undetectable at the end of the cruise to greater than 10 M, and ammonium levels ranged from less than 0.1 to 1.9M. The uptake of both nitrate and ammonium as a function of irradiance was found to be a saturation response. Photoinhibition occurred and was found to be greater for ammonium uptake. Ammonium uptake also saturated at irradiance levels five times lower than those needed to saturate nitrate uptake. Nitrate and ammonium uptake as a function of nitrogen concentration also was characterized by a saturation response, with the estimated half-saturation constant (K _s) value for nitrate uptake being 0.29 M. Elevated ammonium concentrations inhibited nitrate uptake, and the response appeared to be one of exponential decrease with increasing concentrations of ammonium. The most important factor in the Greenland Sea influencing ammonium uptake during the spring was irradiace, while both irradiance and ammonium concentrations played major roles in regulating nitrate uptake and new production.     

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.     

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.     

Desiccation stress in two intertidal beachrock biofilms

Chlorophyll a fluorescence was used to look at the effect of desiccation on the photophysiology in two beachrock microbial biofilms from the intertidal rock platform of Heron Island, Australia. The photophysiological response to desiccation differed between the beachrock microbial communities. The black biofilm from the upper shoreline, dominated by Calothrix sp., showed a response typical of desiccation-tolerant cyanobacteria, where photosynthesis closed down during air exposure with a rapid and complete recovery upon rehydration. In contrast, the pink biofilm from the mid-intertidal zone, dominated by Blennothrix sp., showed no distinct response to desiccation stress and instead maintained reduced photosynthesis throughout drying and re-wetting cycles. Spatial differences in photosynthetic activity within the black biofilm were evident with a faster recovery rate of photosynthesis in the surface cyanobacteria than in the deeper layers of the biofilm. There was no variation with depth in the pink biofilm. The photophysiological differences in desiccation responses between the beachrock biofilms exemplify the ecological niche specialisation of these complex microbial communities, where the functional differences help to explain their vertical distribution on the intertidal shoreline.     

Effect of seawater velocity on inorganic nitrogen uptake by morphologically distinct forms of Macrocystis integrifolia from wave-sheltered and exposed sites

In conditions of low water motion (<0.06 ms^–1), the availability of essential nutrients to macroalgae, and thus their potential productivity, may be limited by thick diffusion boundary-layers at the thallus surface. The ability of macroalgae to take up nutrients in slow moving water may be related to how their blade morphology affects diffusion boundarylayer thickness. For the giant kelp, Macrocystis integrifolia Bory, morphological measurements indicate that blades of plants from a site exposed to wave action are thick, narrow and have a heavily corrugated surface. In contrast, blades from a site with a low degree of water motion are relatively thin, with few surface corrugations and large undulations along their edges. The aim of our work was to test the hypothesis that morphological features of M. integrifolia blades from a sheltered site allow enhanced inorganic nitrogen uptake at low seawater velocities compared to blades with a wave-exposed morphology. The rate of nitrate and ammonium uptake by morphologically distinct blades of M. integrifolia, from sites that were sheltered from and exposed to wave action, were measured in the laboratory at a range of seawater velocities (0.01 to 0.16 ms^–1), between March and May 1993. For both sheltered and exposed blade morphologies, nitrate and ammonium uptake rates increased with increasing seawater velocity, reaching a maximum rate at 0.04 to 0.06 ms^–1. Uptake parameters V _max (maximum uptake rate) and U _0.37 (the velocity at which the uptake rate is 37% of the maximum rate) were estimated using an exponential decay formula. These parameters were similar for both blade morphologies, at all seawater velocities tested. Additional measurements suggest that the nitrogen status of M. integrifolia blades from wavesheltered and exposed sites were similar throughout the experimental period, and thus nitrogen status did not affect the rate of nitrogen uptake in these experiments. on the basis of these results, we conclude that blade morphology does not enhance nitrogen uptake by M. integrifolia in conditions of low water motion. Potential effects of diffusion boundary-layers on kelp productivity are discussed.     

Rehydration of desiccated intertidal brown algae: Release of dissolved organic carbon and water uptake

The release of dissolved organic carbon (DOC) by Ascophyllum nodosum (L.) Le Jol. and Fucus vesiculusus L. during rehydration was investigated after desiccation under experimental conditions. During 30 min of rehydration of plants which had lost up to 70% of their water, A. nodosum released about 2 to 10 mg C/100 g dry weight, while F. vesiculosus released 10 to 50 times more. When dried to water losses exceeding 70%, plants of both species yielded similar amounts of DOC, approximating 2 g C/100 g dry weight. Considerable variance in the amounts of DOC released by individual plant of both species was observed over the whole range of water losses tested. Among the DOC released, the portion of carbohydrate-C was O to 5% with A. nodosum but 2 to 47% with F. vesiculosus. This difference was most pronounced when less than 100 mg C/100 g dry weight were released. A. nodosum regained a smaller percentage of its original weight than F. vesiculosus after 30 min of rehydration when more than 30% of fresh weight were lost during desiccation. Curves were obtained which enable the estimation of water losses in naturally desiccated specimens within 30 min.     

Drought stress,rain and recovery of the intertidal seaweed <Emphasis Type="Italic">Fucus spiralis</Emphasis>

Non-motile organisms of intertidal shores such as seaweeds have to cope with a great variability of environmental factors. In this survey, we studied whether different morphotypes of the intertidal seaweed Fucus spiralis L. are also reflected in a characteristic performance. Desiccation and recovery of this Phaeophyceae were investigated in field experiments near Aljezur, Portugal. Fucus spiralis is exposed to serious desiccation during periods of falling tide, resulting in a tissue water loss of about 90%. Due to large semidiurnal tidal ranges in this area, two morphotypes can be distinguished: F. spiralis growing in the lower intertidal (LZ) is thicker and fleshier compared with plants in the upper intertidal (HZ), and this is reflected in a significant difference in fresh and dry mass. During sunny days and at low tide, effective quantum yields (Φ_PSII) decreased significantly after 2 h desiccation. This continued until re-submersion. The photosynthetic performances of HZ and LZ plants also differed significantly after LZ plants were already submerged and photosynthetisizing, but the HZ specimens still exposed to air. Recovery experiments after desiccation treatments showed fast recovery within 6 min after re-submersion in both morphotypes. HZ specimens showed a slower recovery, which indicates a protection measure to the adverse conditions in the upper intertidal. In 24 h desiccation treatments, however, HZ specimens expressed a significantly higher maximum fluorescence yield F _v /F _m recovery. Simulated rainfalls during low tides caused photosynthetic activity to drop to 50% of initial F _v /F _m , independent of the length of the rain period. Treated plants also fully recovered after 6 min re-submersion in seawater. A comparison of single fronds and tufts clearly indicated advantages of the tuft growth strategy: tufts showed higher Φ_PSII at prolonged emersion times. Our study indicated a clear relationship between size and drought resistance, which was primarily due to the smaller and hardy HZ plants that withstand longer desiccation times without damage.     

Light-dependency of nitrate uptake by phytoplankton over the spring bloom in Auke Bay,Alaska

The effect of light intensity on nitrate uptake by natural populations of phytoplankton was examined by ¹⁵N traceruptake experiments during the spring (March–May 1987) in Auke Bay, Alaska. The data were fit to a rectangular hyperbolic model which included a term for dark uptake. Three types of curves described nitrate uptake as a function of light intensity. The first (Type I) had a low half-saturation light intensity (K _I), low chlorophyll-specific uptakes rates, no dark uptake and occasional photoinhibition. These were observed during a period of biomass decrease, accompanied by low daily light and strong wind, prior to the major bloom. The second type (Type II) had relatively high K _I, high chlorophyll-specific uptake rates, and no dark uptake. Type II curves were observed during most of the period prior to nitrate depletion in the surface waters. Types I and II both appeared prior to nitrate depletion in the water and reflected variations in the light history of the phytoplankton population. The third type (Type III) occurred in nitrate-deplete conditions, when nitrate uptake was less dependent on light intensity (i.e., high rates of dark uptake and lower K _I). Decreased light-dependency during this period was coupled with physiological nitrogen deficiency in the population. Comparing these parameters to those of photosynthetic carbon fixation, K _Ivalues of nitrate uptake were generally higher than those of photosynthesis prior to nitrate depletion, and lower during nutrient-deplete conditions.     

Effects of simulated current on the growth rate and nitrogen metabolism of Gracilaria tikvahiae (Rhodophyta)

In a series of multifactorial laboratory experiments, Gracilaria tikvahiae apical segments were grown in an apparatus in which they were exposed simultaneously to 3 simulated current speeds (7.5, 15, 22.5 cm s^-1) and a still control, and either 3 ammonium concentrations (<6, 37–39, and 119–136 M) under ample uniform light (ca 200 E m^-2 s^-1) or 3 light intensities (ca 35, 90, and 270 E m^-2 s^-1) with uniform surplus ammonium. Growth rates of apical segments were determined in each experiment as well as nitrogen and carbon composition of tissues and fluxes of NH₄, NO₃/NO₂, and PO₄ in media. In a supplementary series of field experiments, apical segments of G. tikvahiae weresimultaneously exposed to 2 different regimes of water motion in adjacent chambers at several sites characterized by widely different ammonium regimes. The application of simulated current significantly enhanced growth rates in all experiments which utilized recently collected plants. Generally, this enhancement was fully realized at 7.5 cm s^-1, with growth rates tending to plateau above that speed. Growth enhancement by simulated current was independent of ammonium concentration and was considerably reduced at the lowest light intensity. In experiments conducted with plants that had been maintained for several months in aquaria, simulated current failed to enhance growth rates. This suggested that growth responses were at least partly a function of prior conditioning. Growth rates were a direct function of light intensity and an inverse function of ammonium concentration, indicating ammonium inhibition at the higher applied concentrations. Simulated current slightly enhanced rates of ammonium uptake but this did not consistently result in reduced C:N ratios, suggesting that the growth-stimulating effect of relative water motion was attributable to factors other than N uptake. There was evidence of luxury consumption of ammonium. In field experiments, growth rates were not significantly related to exposure to water motion.     

Autecology of Fucus distichus ssp. distichus (Phaeophyceae: Fucales) in Nova Scotia,Canada

The autecology of Fucus distichus L. ssp. distichus was investigated at several sites on the Atlantic coast of Nova Scotia, Canada, throughout a 2-year period. This species is confined to high-littoral rock pools at exposed sites, where there appears to be little competition from other organisms. Water temperature, salinity, dissolved oxygen, and hydrogen-ion concentrations underwent both considerable seasonal and short-term variation. Pools with extremes of salinity did not contain F. distichus ssp. distichus. Receptacles are formed during winter, and by late spring they are cast. Sporelings were first visible during September; their development was slow, and these plants did not become reproductive until the second year. During the period February to May, rapid growth of new fronds occurred; an increase in both length and bulk of the plants resulted from this growth. Fronds of mature plants continued to elongate slowly throughout the summer and the early autumn with the formation of receptacles in winter. Plants were frequently damaged, which resulted in regeneration of new fronds from the wounded surface.Issued as NRCC No. 14510