IN SITU UPTAKE KINETICS OF AMMONIUM AND PHOSPHATE AND CHEMICAL COMPOSITION OF THE RED SEAWEED GRACILARIA TIKVAHIAE1

The uptake kinetics of ammonium and phosphate by Gracilaria tikvahiae McLachlan were studied under field conditions. Seaweeds, pulse fed once a week for 6 h over a 4-week period, had maximum uptake rates of 19 μmol·g fwt^?1·h^?1 for ammonium and 0.28 μmol·g fwt^?1·h^?1 for phosphate. For both nutrients there was a positive linear correlation between uptake rate (v) and concentration (S) over the entire range of concentration tested. In a nutrient depletion experiment, the phosphate uptake curve determined over a wide range of concentrations consisted of two stages of saturation at low concentrations, and a linear phase at high concentrations. Ash free dry weight, chlorophyll a, phycoerythrin, and protein content were higher in pulse fed plants than in control plants receiving no nutrient additions, while the reverse held true for carbohydrate contents and the C/N ratios. The C/N ratio inversely correlated with ammonium and phosphate uptake rate as well as protein and phycoerythrin content, and positively with carbohydrate content.     

PHOSPHORUS AND NITROGEN NUTRITION IN CHONDRUS CRISPUS (RHODOPHYTA): EFFECTS ON TOTAL PHOSPHORUS AND NITROGEN CONTENT,CARRAGEENAN PRODUCTION,AND PHOTOSYNTHETIC PIGMENTS AND METABOLISM1

The existence of a phenomenon in phosphorus (P) nutrition comparable to the “Neish effect” in nitrogen (N) nutrition (an inverse relation between seawater N enrichment and carrageenan content) was investigated in the temperate red alga Chondrus crispus Stackhouse. Plants were preconditioned for 17 d and then cultured under varying enrichments of P (0, 3, 6, 10, 15 μM P·wk^?1) and a constant N enrichment (53.5 μM N·wk^?1) for 5 wk. Tissue total P, tissue total N, and carrageenan contents were then determined. Identical experiments were performed using C. crispus collected during the fall, winter, spring, and summer seasons. The procedure was repeated using material collected during the following fall season and cultured under constant P (6 μM P·wk^?1) and varying N enrichments (0, 3, 6, 10, 25 μM N·wk^?1). In the fall (P) experiment, carrageenan content was the highest [53.1 ± 0.3% DW (dry weight)], and tissue total P content was the lowest (1.71 ± 0.27 mg P·g DW^?1) in plants that received no P enrichment. Carrageenan content was stable (46.1 ± 1.8% DW) for plants given enrichments of 3 μM P·wk^?1 and greater. Thus, a decrease in carrageenan content, concomitant with an increase in tissue total P content, was observed, but only at tissue total P levels below 2 mg P·g DW^?1. As these levels were always higher than 2 mg P·g DW^?1 in the winter, spring, and summer experiments, carrageenan content remained constant within each season at 46.2 ± 1.3, 43.1 m 0.7, and 44.5 ± 0.6% DW, respectively. Nitrogen enrichment of plants collected in the fall did not affect carrageenan content, which was stable at 49.3 ± 0.9% DW. When these plants were compared with those of the previous fall experiment (6 μM P·wk^?1 and 53.5 μM N·wk^?1), a slight increase in carrageenan content was noted. Thus, at sufficiently high concentration, N also decreased carrageenan content in C. crispus. Phosphorus nutrition had no significant effect on photosynthesis versus irradiance parameters (P_max, α, R_d, I_c, and I_k), the contents of the photosynthetic pigments chlorophyll-a, phycoerythrin (PE), phycocyanin (PC), and allophycocyanin (APC), and the ratios PE:APC and PC:APC. In contrast, N nutrition affected both P_maxand the photosynthetic pigment contents. The data indicate that N limitation reduces the number of phycobilisomes but not their size. The greater reduction in phycobiliprotein than chlorophyll-acontent corroborates the natural bleaching phenomenon regularly observed in C. crispus populations during summer when N levels are generally low in seawater. These results suggest that C. crispus in the temperate waters of the Bay of Fundy may experience N limitation, but P limitation is unlikely.     

SHIFT FROM CHLOROPHYTES TO CYANOBACTERIA IN BENTHIC MACROALGAE ALONG A GRADIENT OF NITRATE DEPLETION1

A survey of the spatial distribution of benthic macroalgae in a fluvial lake of the St. Lawrence River (Lake Saint‐Pierre, Quebec, Canada) revealed a shift in composition from chlorophytes to cyanobacteria along the flow path of nutrient‐rich waters originating from tributaries draining farmlands. The link between this shift and changes in water quality characteristics was investigated by sampling at 10 sites along a 15 km transect. Conductivity, current, light extinction, total phosphorus (TP; >25 μg P · L^?1), and ammonium (8–21 μg N · L^?1) remained fairly constant along the transect in contrast to nitrate concentrations, which fell sharply. Filamentous and colonial chlorophytes [Cladophora sp. and Hydrodictyon reticulatum (L.) Bory] dominated in the first 5 km where nitrate concentrations were >240 μg N · L^?1. A mixed assemblage of chlorophytes and cyanobacteria characterized a 1 km transition zone where nitrate decreased to 40–80 μg N · L^?1. In the last section of the transect, nitrate concentrations dropped below 10 μg N · L^?1, and cyanobacteria (benthic filamentous mats of Lyngbya wollei Farl. ex Gomont and epiphytic colonies of Gloeotrichia) dominated the benthic community. The predominance of nitrogen‐fixing, potentially toxic cyanobacteria likely resulted from excessive nutrient loads and may affect nutrient and trophic dynamics in the river.     

Zooplankton and bacteria contribution to phosphorus and nitrogen internal cycling in a tropical and eutrophic reservoir: Pampulha Lake,Brazil

Nutrient regeneration and respiration rates of natural zooplankton from a tropical reservoir were experimentally measured. Excretion rates of ammonia (E_a), orthophosphate (E_p) and community respiration rates (R) were estimated considering the variations in the concentrations of ammonia, orthophosphate and dissolved oxygen between control and experimental units. The ranges obtained for these rates from the 2 h assays were E_a = 1.95–4.95 μg N-NH₄ · mg · DW⁻¹ · h⁻¹; E_p = 0.12–0.76 μg P-PO₄ mg DW⁻¹ · h⁻¹. Respiratory rates were quite constant (R = 0.01–0.02 mg O₂ · mg DW⁻¹ · h⁻¹). The uptake of nutrients due to bacteria can affect the experimental determination of excretion rates of zooplankton. Orthophosphate release increased from 0.28 to 0.82 μg P-PO₄ · mg DW⁻¹ · h⁻¹ when bacterial activity was depleted by antibiotic addition in experimental vessels (Exp IV). This demonstrates that free living bacteria are able to consume promptly most phosphorus excreted by zooplankton. Ammonia excretion rates were lower in experimental units containing antibiotics. Lower excretion rates were also obtained with longer exposure times and higher biomass levels in the experimental units. Finally, this study also showed that zooplankton excretion can affect significantly turn over rates of total phosphorus in Pampulha Reservoir. In some periods, specially during the dry season when zooplankton biomass was very high, phosphorus release by zooplankton, during one single day, can be as high as 40% of the total phosphorus content in lake water (Turn over time = 2.5 days).     

OLISTHODISCUS LUTEUS (CHRYSOPHYCEAE). III. UPTAKE AND UTILIZATION OF NITROGEN AND PHOSPHORUS1

Uptake and assimilation of nitrogen and phosphorus were studied in Olisthodiscus luteus Carter. A diel periodicity in nitrate reductase activity was observed in log and stationary phase cultures; there was a 10-fold difference in magnitude between maximum and minimum rates, but other cellular features such as chlorophyll a, carbon, nitrogen, C:N ratio (atoms) · cell^?1 were less variable. K_s values (~2 μM) for uptake of nitrate-N and ammonium-N were observed. Phosphorus assimilated · cell^?1· day^?1 varied with declining external phosphorus concentrations; growth rates <0.5 divisions · day^?1 were common at <0.5 μM PO_4-P. Phosphate uptake rates (K_s= 1.0–1.98 μM) varied with culture age and showed multiphasic kinetic features. Alkaline phosphatase activity was not detected. Comparisons of the nutrient dynamics of O. luteus to other phytoplankton species and the ecological implications as related to the phytoplankton community of Narragansett Bay (Rhode Island) are discussed.     

PHOTOSYNTHESIS AND RESPIRATION OF THE CONCHOCELIS STAGE OF ALASKAN PORPHYRA (BANGIALES,RHODOPHYTA) SPECIES IN RESPONSE TO ENVIRONMENTAL VARIABLES1

Photosynthesis and respiration of three Alaskan Porphyra species, P. abbottiae V. Krishnam., P. pseudolinearis Ueda species complex (identified as P. “pseudolinearis” below), and P. torta V. Krishnam., were investigated under a range of environmental parameters. Photosynthesis versus irradiance (P–I) curves revealed that maximal photosynthesis (P_max), irradiance at maximal photosynthesis (I_max), and compensation irradiance (I_c) varied with salinity, temperature, and species. The P_max of Porphyra abbottiae conchocelis varied between 83 and 240 μmol O₂ · g dwt^?1 · h^?1 (where dwt indicates dry weight) at 30–140 μmol photons · m^?2 · s^?1 (I_max) depending on temperature. Higher irradiances resulted in photoinhibition. Maximal photosynthesis of the conchocelis of P. abbottiae occurred at 11°C, 60 μmol photons · m^?2·s^?1, and 30 psu (practical salinity units). The conchocelis of P. “pseudolinearis” and P. torta had similar P_max values but higher I_max values than those of P. abbottiae. The P_max of P. “pseudolinearis” conchocelis was 200–240 μmol O₂ · g dwt^?1 · h^?1 and for P. torta was 90–240 μmol O₂ · g dwt^?1 · h^?1. Maximal photosynthesis for P. “pseudolinearis” occurred at 7°C and 250 μmol photons · m^?2 · s^?1 at 30 psu, but P_max did not change much with temperature. Maximal photosynthesis for P. torta occurred at 15°C, 200 μmol photons · m^?2 · s^?1, and 30 psu. Photosynthesis rates for all species declined at salinities <25 or >35 psu. Estimated compensation irradiances (I_c) were relatively low (3–5 μmol · photons · m^?2 · s^?1) for intertidal macrophytes. Porphyra conchocelis had lower respiration rates at 7°C than at 11°C or 15°C. All three species exhibited minimal respiration rates at salinities between 25 and 35 psu.     

KINETICS OF SILICIC ACID UPTAKE AND RATES OF SILICA DISSOLUTION IN THE MARINE DIATOM THALASSIOSIRA PSEUDONANA1,2

Tracer techniques using the stable isotope ³⁰Si were used to measure rates of silicic acid uptake and silica dissolution in silicon replete and silicon depleted populations of 2 clones of the marine diatom Thalassiosira pseudonana Hasle & Heimdal. Uptake kinetics were describable using the Michaelis-Menten equation for enzyme kinetics, and no threshold concentration for uptake was evident. The maximum specific uptake rate of the estuarine clone 3H (0.062–0.092 · h^?1) was higher than that of the Sargasso Sea clone 13-1 (0.028–0.031 · h^?1), but half-saturation constants for uptake by the 2 clones were not measurably different (0.8–2.3 μM for 3H; 1.4–1.5 μM for 13-1). There was little or no light dependence of uptake in populations grown under optimal light conditions prior to the experiment. Exponentially growing populations released silicic acid to the medium by dissolution of cellular silica at rates ranging from 6.5 to 15% of the maximum uptake rate.     

SEASONAL PHOTOSYNTHETIC PERFORMANCE OF MACROCYSTIS INTEGRIFOLIA (PHAEOPHYCEAE)1

The seasonal photosynthetic performances of three age classes of blades of Macrocystis integrifolia Bory were estimated by studying their photosynthetic rate vs. irradiance curves and pigment contents for 15 months. All blade types were irradiance-saturated between 25 and 70 μE · m^?2· S^?1. Young and mature blade tissues had higher photosynthetic maxima and initial slopes on an area basis than older blade tissue. The latter, however, had pigment concentrations similar to those in mature blade tissues. All these parameters varied on a seasonal basis. The photosynthetic maxima ranged from 0.1–0.8 μmol · C · cm^?2· h^?t and showed two peaks, one in late summer-early fall and the other in late winter. Changes in initial slope and pigment concentrations in the blade tissues suggest that, changes in the size or efficiency of electron transfer in the photosynthetic unit occur. These data are discussed in relation to changes in seawater temperature and nitrate concentrations.     

RAPID NITRATE UPTAKE RATES AND LARGE SHORT‐TERM STORAGE CAPACITIES MAY EXPLAIN WHY OPPORTUNISTIC GREEN MACROALGAE DOMINATE SHALLOW EUTROPHIC ESTUARIES1

We quantified the effects of initial macroalgal tissue nitrogen (N) status (depleted and enriched) and varying pulses of nitrate (NO₃^?) concentration on uptake and storage of nitrogen in Ulva intestinalis L. and Ulva expansa (Setch.) Setch. et N. L. Gardner using mesocosms modeling shallow coastal estuaries in Mediterranean climates. Uptake of NO₃^? (μmol · g dry weight [dwt]^?1 · h^?1) was measured as loss from the water after 1, 2, 4, 8, 12, and 24 h and storage as total tissue nitrogen (% dwt) and nitrate (ppm). Both species of algae exhibited a high affinity for NO₃^? across all N pulses and initial tissue contents. There was greater NO₃^? removal from the water for depleted than enriched algae across all time intervals. In the low‐N‐pulse treatment, U. intestinalis and U. expansa removed all measurable NO₃^? within 8 and 12 h, respectively, and in the medium and high treatments, removal was high and then decreased over time. Maximum mean uptake rates of nitrate were greater for U. expansa (～300 μmol · g dwt^?1 · h^?1) than U. intestinalis (～100 μmol · g dwt^?1 · h^?1); however, uptake rates were highly variable over time. Overall, U. expansa uptake rates were double those of U. intestinalis. Maximum tissue NO₃^? for U. expansa was >1,000 ppm, five times that of U. intestinalis, suggesting that U. expansa has a greater storage capacity in this cellular pool. These results showed that opportunistic green algae with differing tissue nutrient histories were able to efficiently remove nitrate from the water across a wide range of N pulses; thus, both are highly adapted to proliferate in estuarine environments with pulsed nutrient supplies.     

Seasonal nitrate physiology of Macrocystis integrifolia Bory

Ambient sea-water nitrate and tissue nitrogen (ethanol soluble nitrate and amino acids, as well as total nitrogen) of Macrocystis integrifolia Bory were monitored over a 2-yr period in Bamfield, Vancouver Island, British Columbia. Sea-water nitrate varied from a high of 12 μmol · 1^?1 (individual values as high as 23 μmol · 1^?1 were recorded) in late winter to below detection limits for most of the summer. Tissue nitrate and total nitrogen paralleled the ambient nitrate levels and showed summer minima and winter maxima (from 0 to 70 μmol · g fresh wt^?1 for nitrate and from 0.8 to 2.9% of dry wt for total N). The nitrate uptake capacity was inversely proportional to tissue nitrate concentration and, furthermore, was much higher for subapical surface blades (60–70 nmol · cm^?2 · h^?1) than for older, deeper blades (5–10 nmol · cm^?2 · h^?1). Nitrate uptake by subapical blade disks in summer is apparently higher in dark (1.0–1.7 μmol · g fresh wt^?1 · h^?1) than in light (0.6–1.3 μmol · g fresh wt^?1 · h^?1) and the data obtained in 36–108 h experiments indicate nitrate pool sizes of 30–90 μmol · g fresh wt^?1. These pools are $\text{2}\text{3}$ to nearly full in winter. Ammonium does not inhibit nitrate uptake. It is taken up and apparently utilized much faster than nitrate and it may well be an important source of nitrogen for marine macrophytes.     

PRODUCTION AND NUTRIENT REMOVAL BY PERIPHYTON GROWN UNDER DIFFERENT LOADING RATES OF ANAEROBICALLY DIGESTED FLUSHED DAIRY MANURE1

Growing algae to scrub nutrients from manure presents an alternative to the current practice of land application and provides utilizable algal biomass as an end product. The objective of this study was to assess algal growth, nutrient removal, and nitrification using higher light intensities and manure loading rates than in the previous experiments. Algal turfs, with periphyton mainly composed of green algal species, were grown under two light regimes (270 and 390 μmol photons·m^?2· s^?1) and anaerobically digested flushed dairy manure wastewater (ADFDMW) loading rates ranging from 0.8 to 3.7 g total N and 0.12 to 0.58 g total P·m^?2·d^?1. Filamentous cyanobacteria (Oscillatoria spp.) and diatoms (Navicula, Nitzschia, and Cyclotella sp.) partially replaced the filamentous green algae at relatively high ADFDMW loading rates and more prominently under low incident light. Mean algal production increased with loading rate and irradiance from 7.6±2.71 to 19.1±2.73 g dry weight· m^?2·d^?1. The N and P content of algal biomass generally increased with loading rate and ranged from 2.9%–7.3% and 0.5%–1.3% (by weight), respectively. Carbon content remained relatively constant at all loading rates (42%–47%). The maximum removal rates of N and P per unit algal biomass were 70 and 13 mg·g^?1 dry weight·m^?2·d^?1, respectively. Recovery of nutrients in harvested algal biomass accounted for about 31%–52% for N and 30%–59% for P. Recovery of P appeared to be uncoupled with N at higher loading rates, suggesting that algal potential for accumulation of P may have already been saturated. It appears that higher irradiance level enhancing algal growth was the overriding factor in controlling nitrification in the algal turf scrubber units.     

PRODUCTIVITY OF THE FILAMENTOUS ALGA PITHOPHORA OEDOGONIA (CHLOROPHYTA) IN SURREY LAKE,INDIANA1

Biomass, akinete numbers, net photosynthesis, and respiration of Pithophora oedogonia were monitored over two growing seasons in shallow Surrey Lake, Indiana. Low rates of photosynthesis occurred from late fall to early spring and increased to maximum levels in late spring to summer (29–39 mgO₂·g^?1 dry wt·h^?1). Areal biomass increased following the rise in photosynthesis and peaked in autumn (163–206g dry wt·m^?2). Photosynthetic rates were directly correlated with temperature, nitrogen, and phosphorus over the entire annual cycle and during the growing season. Differences in photosynthetic activity and biomass between the two growing seasons (1980 and 1981) were apparently related to higher, early spring temperatures and higher levels of NO₃-N and PO₄-P in 1981. Laboratory investigations of temperature and light effects on Pithophora photosynthesis and respiration indicated that these processes were severely inhibited below 15°C. The highest P_max value occurred at 35°C (0.602 μmol O₂·mg^?1 chl a·min^?1). Rates of dark respiration did not increase above 25°C thus contributing to a favorable balance of photosynthetic production to respiratory utilization at high temperatures. Light was most efficiently utilized at 15°C as indicated by minimum values of I_k(47 μE·m^?2·s^?1) and I_c (6 μE·m^?2·s^?1). Comparison of P. oedogonia and Cladophora glomerata indicated that the former was more tolerant of temperatures above 30°C. Pithophora's tolerance of high temperature and efficient use of low light intensity appear to be adaptive to conditions found within the dense, floating algal mats and the shallow littoral areas inhabited by this filamentous alga.     

NUTRIENT REQUIREMENTS OF THE DINOFLAGELLATE PERIDINIUM GATUNENSE1

Optimum nutrient conditions for growth and photosynthesis of Peridinium gatunense (Nygaard) (Peridinium cinctum fa. westii) were investigated using axenic clones in batch cultures. Selenium (Se) had previously been found to be an indispensable growth factor for P. gatunense. Optimal, suboptimal, and supraoptimal concentrations of HCO₃^?, N, Ca, Cl, Mg, P, K, S, Si, EDTA-Na, Fe, Mo, Zn, Mn, Co, Se, B, Br, I, and various trace element mixtures were determined by measuring biomass development, growth rates, ¹⁴C uptake, and/or oxygen production at various concentration gradients of these elements. The general characteristics of the best formulation, medium-L 16, relative to other media, are its high content of NaHCO₃ (1 meq · L^?1) and Mo (0.2 μM) but low concentrations of NO_3-N (150 μM), PO_4-P (10 μM), and Fe (0.4 μM), in addition to its content of Se. The total content of trace metals, except for Se, may be reduced to one-fourth of that in medium-L 16 without altering the major growth-promoting properties of the medium. Medium-L 16 deviated considerably from Lake Kinneret (Israel) water, being much lower in macroelements except for N and P. The pH (8.1–8.4) was in the same range, but the values of conductivity (140 μS · cm^?1), alkalinity (1 meq · L^?1) and NaCl (200 μM) were > 8, 2, and 30 times higher, respectively, in the lake water. Selenium deficiency may limit the growth of P. gatunense in this lake.     

THE RELATIONSHIP OF LIPIDS WITH LIGHT AND CHLOROPHYLL MEASUREMENTS IN FRESHWATER ALGAE AND PERIPHYTON1

Lipid content and lipid class composition were determined in stream periphyton and the filamentous green algae Cladophora sp. and Spirogyra sp, Sterols and phospholipids were compared to chlorophyll a (chl a) as predictors of biomass for stream periphyton and algae. Chlorophyll a, phospholipids, and sterols were each highly correlated with ash-free dry mass (AFDM) (r² > 0.98). Stream periphyton exposed naturally to high light (HL) and low light (LL) had chl a concentrations (μg chl a-mg^?1AFDM) of 7.9± 0.7 and 12.4 ± 2.9, respectively, while the sterol concentrations of these HL and LL stream periphyton (1.6 ± 0.4) were not significantly different (P > 0.05). Periphyton exposed to an irradiance of 300 μmol photons·m^?2s^?1 in the laboratory for 60 h had 5.6 ± 0.55 μg chl a·mg^?1 AFDM, but the same periphyton exposed to 2% incident light for the same amount of time had 11.0 ± 0.56 μg chl a·mg^?1 AFDM. Sterol concentrations in these periphyton communities remained unchanged (1.5 ± 0.3 μg·mg^?1AFDM), Similar results (i.e. changes in chl a but stability of sterol concentrations in response to irradiance changes) were also found for Cladophora and Spirogyra in laboratory experiments. Sterols can be quantified rapidly from a few milligrams of algae and appear to be a useful predictor of eukaryote biomass, whereas cellular levels of chl a vary substantially with light conditions. Phospholipids (or phospholipid fatty acids) are considered to be a reliable measure of viable microbial biomass. Nevertheless, phospholipid content varied substantially and unpredictably among algae and periphyton under different light regimes. Irradiance also had a significant effect on storage lipids: HL Cladophora and HL periphyton had 2 × and 5 × greater concentrations of triacylglycerols, respectively, compared to their LL forms. HL and LL algae also differed in the concentration of several major fatty acids. These light-induced changes in algal lipids and fatty acids have important implications for grazers.     

CELL‐SPECIFIC EXTRACELLULAR PHOSPHATASE ACTIVITY OF DINOFLAGELLATE POPULATIONS IN ACIDIFIED MOUNTAIN LAKES1

High bulk extracellular phosphatase activity (PA) suggested severe phosphorus (P) deficiency in plankton of three acidified mountain lakes in the Bohemian Forest. Bioavailability of P substantially differed among the lakes due to differences in their P loading, as well as in concentrations of aluminum (Al) and its species, and was accompanied by species‐specific responses of phytoplankton. We combined the fluorescently labeled enzyme activity (FLEA) assay with image cytometry to measure cell‐specific PA in natural populations of three dinophyte species, occurring in all the lakes throughout May–September 2007. The mean cell‐specific PA varied among the lakes within one order of magnitude: 188–1,831 fmol · cell^?1 · h^?1 for Gymnodinium uberrimum (G. F. Allman) Kof. et Swezy, 21–150 fmol · cell^?1 · h^?1 for Gymnodinium sp., and 22–365 fmol · cell^?1 · h^?1 for Peridinium umbonatum F. Stein. To better compare cell‐specific PA among the species of different size, the values were normalized per unit of cell biovolume (amol · μm^?3 · h^?1) for further statistical analysis. A step‐forward selection identified concentrations of total and ionic Al together with pH as significant factors (P < 0.05, Monte Carlo permutation test), explaining cumulatively 57% of the total variability in cell‐specific PA. However, this cell‐specific PA showed an unexpected reverse trend compared to an overall gradient in P deficiency of the lake plankton. The autecological insight into dinophyte cell‐specific PA therefore suggested other factors, such as light availability, mixotrophy, and/or zooplankton grazing, causing further PA variations among the acidified lakes.     

Nutrient uptake and alkaline phosphatase (ec 3:1:3:1) activity of emiliania huxleyi (PRYMNESIOPHYCEAE) during growth under n and p limitation in continuous cultures

Emiliania huxleyi (strain L) expressed an exceptional P assimilation capability. Under P limitation, the minimum cell P content was 2.6 fmol P·cell^?1, and cell N remained constant at all growth rates at 100 fmol N·cell^?1. Both, calcification of cells and the induction of the phosphate uptake system were inversely correlated with growth rate. The highest (cellular P based) maximum phosphate uptake rate (V_max^P) was 1400 times (i.e. 8.9 h^?1) higher than the actual uptake rate. The affinity of the P‐uptake system (dV/dS) was 19.8 L·μmol^?1·h^?1 at μ = 0.14 d^?1. This is the highest value ever reported for a phytoplankton species. V_max and dV/dS for phosphate uptake were 48% and 15% lower in the dark than in the light at the lowest growth rates. The half‐saturation constant for growth was 1.1 nM. The coefficient for luxury phosphate uptake (Q_maxt/Q_min) was 31. Under P limitation, E. huxleyi expressed two different types of alkaline phosphatase (APase) enzyme kinetics. One type was synthesized constitutively and possessed a V_max and half‐saturation constant of 43 fmol MFP·cell^?1·h^?1 and 1.9 μM, respectively. The other, inducible type of APase expressed its highest activity at the lowest growth rates, with a V_max and half‐saturation constant of 190 fmol MFP·cell^?1·h^?1 and 12.2 μM, respectively. Both APase systems were located in a lipid membrane close to the cell wall. Under N‐limiting growth conditions, the minimum N quotum was 43 fmol N·cell^?1. The highest value for the cell N‐specific maximum nitrate uptake rate (V_max^N) was 0.075 h^?1; for the affinity of nitrate uptake, 0.37 L·μmol^?1·h^?1. The uptake rate of nitrate in the dark was 70% lower than in the light. N‐limited cells were smaller than P‐limited cells and contained 50% less organic and inorganic carbon. In comparison with other algae, E. huxleyi is a poor competitor for nitrate under N limitation. As a consequence of its high affinity for inorganic phosphate, and the presence of two different types of APase in terms of kinetics, E. huxleyi is expected to perform well in P‐controlled ecosystems.     

PHYSIOLOGICAL VARIATION AND ELECTROPHORETIC BANDING PATTERNS OF GENETICALLY DIFFERENT SEASONAL POPULATIONS OF SKELETONEMA COSTATUM (BACILLARIOPHYCEAE)1

Clones of Skeletonema costatum (Grev.) Cl. isolated from Narragansett Bay, R.I., during different seasons were grouped according to their electrophoretic banding patterns. The growth rates, pg chlorophyll · cell^?1, carbon uptake · cell^?1· h^?1, and carbon uptake · pg chl^?1· h^?1 were measured at 20°C, in a 14:10 h L:D cycle at 180 μE · m^?2· s^?1. Statistically significant sources of variation were found among groups of clones in growth rate, pg chl · cell^?1, and carbon uptake · pg chl^?1· h^?1. It was concluded that there is a significant relationship between the physiological characteristics of clones isolated from populations in different seasons and patterns of genetic variation inferred from the electrophoretic studies. However, genetic diversity detected by banding patterns tends to underestimate the total genetic diversity in natural populations. The groups of clones most common in summer bloom populations had significantly higher growth rates, lower values of pg chl · cell^?1, and higher rates of carbon uptake · pg chl^?1· h^?1 at 20°C than did the group of clones most common in winter bloom populations. However, differences among groups in these parameters at 20°C alone cannot account for the seasonal cycling of genetically variable populations of Skeletonema in Narragansett Bay. The range of growth rates among clones of this species is 0.1–5.0 divisions · d^?1 under a single set of temperature and light conditions. Chlorophyll concentrations range from 0.2–1.7 pg chl · cell^?1 and carbon uptake · pg chl^?1· h^?1 varies by a factor of 7 among clones. The range of physiological variation in this species means that it is difficult to use laboratory studies of single clones to analyze the responses of natural populations of Skeletonema.     

THE UPTAKE OF IODATE BY MARINE PHYTOPLANKTON1

Several studies have suggested that phytoplankton play a role in the iodine cycle. Using a short-term incubation technique for determining the uptake of iodate by phytoplankton, cultures of Thalassiosira oceanica Hasle, Skeletonema costatum (Greville) Cleve, Emiliania huxleyi (Lohmann) Hay and Mohler, and Dunaliella tertiolecta Butcher were found to be capable of assimilating iodate at rates ranging from 0.003 to 0.24 nmol IO₃^?·μg chlorophyll a^?1·h^?1. The kinetics for the uptake of iodate can be modeled, and the similarity between the model and experimental results suggests that there is a steady state between iodate uptake and release of dissolved iodine from the cells, presumably in the form of iodide. Two experiments were conducted in the Sand Shoal Inlet of the Cobb Bay estuary (37°15′N, 75°50′W). The uptake of iodate was 0.26 and 0.08 nmol IO₃^?·μg chlorophyll a^?1·h^?1 during high and low tide, respectively. Using field estimates based on measured levels of iodate in the estuary, we estimate that phytoplankton can take up as much as 3% of the ambient pool of iodate on a daily basis and the entire pool in about 1 month. Thus, phytoplankton can be a significant component of the global iodine cycle by mediating changes in the speciation of iodine in the marine environment.     

PHYCOBILIN CONTENT OF THE CONCHOCELIS PHASE OF ALASKAN PORPHYRA (BANGIALES,RHODOPHYTA) SPECIES: RESPONSES TO ENVIRONMENTAL VARIABLES1

Variations of pigment content in the microscopic conchocelis stage of four Alaskan Porphyra species were investigated in response to environmental variables. Conchocelis filaments were cultured under varying conditions of irradiance and nutrient concentrations for up to 60 d at 11°C and 30 psu salinity. Results indicate that conchocelis filaments contain relatively high concentrations of phycobilins under optimal culture conditions. Phycobilin pigment production was significantly affected by irradiance, nutrient concentration, and culture duration. For Porphyra abbottiae V. Krishnam., Porphyra sp., and Porphyra torta V. Krishnam., maximal phycoerythrin (63.2–95.1 mg · g dwt^?1) and phycocyanin (28.8–64.8 mg · g dwt^?1) content generally occurred at 10 μmol photons · m^?2 · s^?1, f/4–f/2 nutrient concentration after 10–20 d of culture. Whereas for Porphyra hiberna S. C. Lindstrom et K. M. Cole, the highest phycoerythrin (73.3 mg · g dwt^?1) and phycocyanin (70.2 mg · g dwt^?1) content occurred at 10 μmol photons · m^?2 · s^?1, f nutrient concentration after 60 d in culture. Under similar conditions, the different species showed significant differences in pigment content. P. abbottiae had higher phycoerythrin content than the other three species, and P. hiberna had the highest phycocyanin content. P. torta had the lowest phycobilin content.     

UPTAKE AND RELEASE OF NITROGEN BY THE MACROALGAE GRACILARIA VERMICULOPHYLLA (RHODOPHYTA)1

Macroalgae, often the dominant primary producers in shallow estuaries, can be important regulators of nitrogen (N) cycling. Like phytoplankton, actively growing macroalgae release N to the water column; yet little is known about the quantity or nature of this release. Using ¹⁵N labeling in laboratory and field experiments, we estimated the quantity of N released relative to assimilation and gross uptake by Gracilaria vermiculophylla (Ohmi) Papenfuss (Rhodophyta, Gracilariales), a non‐native macroalgae. Field experiments were carried out in Hog Island Bay, a shallow back‐barrier lagoon on the Virginia coast where G. vermiculophylla makes up 85%–90% of the biomass. There was good agreement between laboratory and field measurements of N uptake and release. Daily N assimilation in field experiments (32.3±7.2 μ mol N·g dw^?1·d^?1) was correlated with seasonal and local N availability. The average rate of N release across all sites and dates (65.8±11.6 μ mol N·g dw^?1·d^?1) was 67% of gross daily uptake, and also varied among sites and seasons (range=33%–99%). Release was highest when growth rates and nutrient availability were low, possibly due to senescence during these periods. During summer biomass peaks, estimated N release from macroalgal mats was as high as 17 mmol N·m^?2·d^?1. Our results suggest that most estimates of macroalgal N uptake severely underestimate gross N uptake and that N is taken up, transformed, and released to the water column on short time scales (minutes–hours).