Regulation of growth in Laminaria digitata: use of in-vivo nitrate reductase activities as an indicator of nitrogen limitation in field populations of Laminaria spp.

The seasonal variation in growth rate of a population of Laminaria digitata (Huds.) Lamour growing at Arbroath, Scotland was studied between August 1981 and September 1982, and was found to follow the biphasic annual cycle typical of this genus. Growth rates were maximum (0.3 cm cm^-1 mo^-1) in early June and minimum (0.05 cm cm^-1 mo^-1) between September and January. An analysis of the relationship between the seasonal changes in environmental factors (inorganic nitrogen concentrations, irradiance and temperature) with those of growth rate and the accumulation or mobilisation of cellular reserves of carbohydrates and nitrate, indicated that growth was nitrogen-limited between June and October and light-limited (with a possible co-involvement of temperature) for the remainder of the year. These conclusions were supported by the seasonal changes in the ratio of actual: potential in-vivo nitrate reductase activities in L. digitata, thus confirming the suitability of this technique for monitoring the occurrence of nitrogen limitation in Laminaria spp. The seasonal changes in blade nitrate reductase activities closely followed those of growth rate, with maximum activities [0.3 mol NO ₃ ^- reduced g^-1 (wet wt) h^-1] being present in late May and minimum levels [0.01 mol NO ₃ ^- reduced g^-1 (wet wt) h^-1] occurring between November and March. The correlation observed between nitrate reductase activities and growth rate is consistent with the ability of Laminaria spp. to store excess inorganic nitrogen, available during winter and early spring, as NO ₃ ^- , and with the requirement to conserve enzyme protein during the summer period of nitrogen limitation.     

Phosphorus- and nitrogen-limited photosynthesis and growth of Gracilaria tikvahiae (Rhodophyceae) in the Florida Keys: an experimental field study

The relative effects of NH ₄ ⁺ (N) and PO ₄ ^3- (P) on growth rate, photosynthetic capacity (P_max), and levels of chemical constituents of the red macroalga Gracilaria tikvahiae McLachlan were assayed during winter and summer, 1983 in inshore waters of the Florida Keys by using in-situ cage cultures. During winter, both N and P enrichment enhanced growth over that of ambient seawater; however, P rather than N accounted for more (60%) of the increased winter growth. During summer, P, but not N, enhanced growth over ambient seawater and accounted for 80% of increased growth. Similarly, P_max was enhanced by both P and N during winter (but mostly by P) and only by P during summer. Elevated C:P, C:N and N:P ratios of G. tikvahiae tissue during winter, but only C:P and N:P ratios during summer, support the pattern of winter N and P limitation and summer P-limitation. This seasonal pattern of N vs P limited growth of G. tikvahiae appears to be a response to seasonally variable dissolved inorganic N (twofold greater concentrations of NH ₄ ⁺ and NO ₃ ^- during summer compared to winter) and constantly low to undetectable concentrations of PO ₄ ^3- . Mean C:P and N:P ratios of G. tikvahiae tissue during the study were 1 818 and 124, respectively, values among the highest reported for macroalgae.     

Nitrogen exchange between a southeastern USA salt marsh ecosystem and the coastal ocean

The salt marsh ecosystem at North Inlet, South Carolina, USA consistently exported dissolved inorganic nitrogen via tidal exchange with the coastal Atlantic Ocean. Concentrations centrations of NH ₄ ⁺ and NO ₃ ^- +NO ₂ ^- displayed distinct tidal patterns with rising values during ebb flow. These patterns suggest the importance of biogeochemical processes in the flux of material from the salt marsh. NH ₄ ⁺ export peaked during the summer (15 to 20 mg m^-2 tide^-1) during a net balance of tidal water exchange. Remineralization of NH ₄ ⁺ within the salt marsh system appears to be contributing to the estimated annual net export of bout 4.7 g NH ₄ ⁺ -N m^-2 yr^-1. NO ₃ ^- +NO ₂ ^- exports were higher in the fall and winter of 1979 (2 to 4 mg N m^-2 tide^-1). The winter export coincided with a considerable net export of water with no distinctive concentration patterns, suggesting a simple advective export. However, the fall peak of NO ₃ ^- +NO ₂ ^- export occurred during a period of net water balance in tidal exchange and an insignificant freshwater input from the western, forested boundary. During the summer and fall, tidal concentration patterns were particularly apparent, suggesting that nitrification within the salt marsh system was contributing to the estimated annual net export of ca 0.6 g NO ₃ ^- +NO ₂ ^- -N m^-2 yr^-1.Contribution No. 637 from the Belle W. Baruch Institute of Marine Biology and Coastal Research     

Composition chimique élémentaire (C,H, N) et équivalent énergie d'Acartia clausi (Crustacea: Copepoda), espèce importante dans la bioénergétique des écosystèmes côtiers de Méditerranée nord-occidentale

Fixed-point sampling of a shallow tidal estuary was performed hourly for 14 d in summer of 1982 and again in winter of 1983. This sampling regime was of statistically appropriate duration to allow characterization of the variability between periods of 2 to 96 h by spectral analysis of the time-series. The project (PULSE) took place in the Newport River Estuary, located behind the Outer Banks of North Carolina, USA. In all, twentyeight parameters were monitored, encompassing the meteorology, hydrology, water chemistry and phytoplankton-production physiology. Although the annual cycle was monitored, only the winter and summer seasons are compared here, i.e., the lowest water temperatures with the highest water temperatures. The physics, chemistry and biology of the estuary at the hourly scale were highly variable and non-random. The estuary is riverine in winter; growth-limiting nitrogen is supplied as nitrate (NO ₃ ^- ) and ammonium (NH ₄ ⁺ ) by runoff from the drainage basin. In summer, the estuary is lagoonal; nitrogen is supplied as NH ₄ ⁺ by biological regeneration. Chlorophyll a biomass varies primarily at the 4 d period in winter and at the diel period in summer. Although finely tuned to environmental variability, phytoplankton abundances were at equilibrium insofar as daily chlorophyll production was balanced by losses, i.e., grazing, export and deposition. Most important, high-frequency processes, here periods at the scale of cell-division times, can be very important in phytoplankton ecology.     

Seasonal patterns of growth and storage in Laminaria longicruris in relation to differing patterns of availability of nitrogen in the water

The annual cycles of growth and of internal storage of nitrogen and carbon in Laminaria longicruris were compared at three sites in Nova Scotia, Canada. One site had abundant nitrogen all year round, on account of local upwelling, one had abundant nitrogen for only four months and the last was intermediate in this respect. Where nitrogen was abundant year round, growth followed the seasonal pattern of light, and there was little storage of either carbon or inorganic nitrogen. Where nitrogen was only abundant in the sea in winter, growth accelerated at this time and continued throught early summer while internal reserves of nitrogen were utilized. Carbon was stored in summer and used for growth in the fall. At the intermediate site, larger reserves of nitrogen were built up in winter and a high growth rate was maintained through most of the summer. Smaller carbon reserves were accumulated and growth was minimal in winter. Experiments in which plants were transplanted between sites were not conclusive, but suggested that plants at different sites may have genetic differences.     

Seasonal variation in total and soluble tissue nitrogen of Pleurophycus gardneri (Phaeophyceae: Laminariales) in relation to environmental nitrate

Seasonal variations in tissue nitrogen (ethanol soluble nitrate and ninhydrin positive substances, as well as total nitrogen) of different thallus parts of Pleurophycus gardneri Setchell and Saunders were monitored simultaneously with ambient seawater nitrate from 1982 until 1984 in Bamfield, Vancouver Island, British Columbia, Canada. A trend of low, nearly zero levels in ambient nitrate typical for the area in late spring and early summer normally contrasts with average nitrate concentrations of 10 mol NO₃ ^- l^-1 in late fall and winter. Total nitrogen content was greater in the perennial thallus parts, stipe and holdfast than in the annual blade and peaked in fall and early winter. The longitudinal thallus distribution of nitrate revealed a distinct and significant concentration of nitrate in the haptera reaching at maximum 8% nitrate-N of the internal total nitrogen. Internal nitrate concentration ranged from 20 to 5 000 times the ambient nitrate concentration in the midrib, and from 40 to 3 100 times in the wing, while the range was greatest with 400 to 14 000 times in the haptera. P. gardneri contained at most about 7 mol NO₃ ^- g fresh wt^-1 in the blade, which corresponds to about 6% of total tissue nitrogen. Ninhydrin positive substances comprised the major portion of the soluble N pool in P. gardneri and showed a pronounced seasonality. Concentrations of ninhydrin positive substances ranged from 20 to 800 g N g fresh wt^-1 in the midrib and in the wing. In the stipe, ninhydrin positive substances varied from 180 to 2 200 g N g fresh wt^-1, and from 250 to 1 200 g N g fresh wt^-1 in the haptera. Evidence is given that (1) the perennial parts, stipe and haptera of P. gardneri contain the majority of nitrogen products independent of season and ontogenetic stage; (2) ninhydrin positive substances are the most abundant internal nitrogen constituents; (3) the low N values in the blade in summer suggest a nitrogen limited growth; and (4) nitrate may not be the predominant external nitrogen source.     

Seasonal growth and composition of fronds of Macrocystis pyrifera in the Falkland Islands

Frond growth of Macrocystis pyrifera in the Falkland Islands was monitored in shallow coastal water from December 1985 to March 1987, and at a different site in deeper water from December 1985 to June 1986. Growth rates in the deeper bed were generally higher than those recorded in the coastal zone. At both sites, node initiation and elongation rate fluctuated according to the seasonal pattern of light or water temperature. In the shallow coastal area, nitrate was abundant in the winter and below detection levels during late spring and summer. Correlation analysis suggests that the production of the fronds of the giant kelp in this area was probably inhibited during the summer months by extremely low concentrations of nutrients. Internal nitrogen was exhausted approximately one month after a sharp decline in ambient nitrate concentration, and carbon reserves were formed. In the deeper bed of M. pyrifera, nitrogen was abundant all year round and the production of the fronds reflected the seasonal pattern of light or water temperature. The nitrogen content of the tissue probably did not drop below a level that limited production, and no internal carbon reserves were accumulated.     

Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera in a low-nitrogen environment

An adult giant kelp plant (Macrocystis pyrifera), moved from an inshore kelp forest to an offshore, low-nitrogen environment near Santa Catalina Island, California (USA), maintained growth for 2 wk on internal nitrogen reserves. Frond elongation rates decreased significantly during the third week, and plant growth rate (wet wt) dropped from an initial inshore rate of 3.6 to 0.9% d^-1. During this 3 wk period, nitrogen contents and free amino acid concentrations decreased, while mannitol and dry contents increased in frond tissues. After depletion of internal nitrogen reserves, the nitrogen content of lamina and stipe tissues averaged 1.1 and 0.7% dry wt, respectively. The experimental plant was exposed to higher ambient nitrogen concentrations during the last 2 wk. Rates of frond elongation and plant growth increased, but nitrogen content and amino acids in frond tissues remained low. Of the total nitrogen contained in frond tissue of the plant before transplantation, 58% was used to support growth in the absence of significant external nitrogen supply. Amino acids constituted a small proportion of these internal nitrogen reserves. Net movement of nitrogen occurred within large fronds, but not between different frond size classes. The nitrogen content of holdfast tissue remained relatively constant at 2.4% dry wt and accounted for 18 to 29% of the total nitrogen. Holdfast nitrogen was not used to support growth of nitrogen-depleted fronds. In comparison to Laminaria longicruris, which is adapted to long seasonal periods of low nitrogen availability, M. pyrifera has small nitrogen-storage capacity. However, internal reserves of M. pyrifera appear adequate to make nitrogen starvation uncommon in southern California kelp forests.     

Ammonium regeneration by microzooplankton in the Oslofjord

Concentrations of dissolved inorganic nitrogen compounds above the pycnocline in the Oslofjord are very low in the summer, with turnover times of the inorganic N pools of no more than a few hours. To investigate the possibility that continued phytoplankton growth in the summer depends on ammonium excretion by microzooplankton, rates of NH ₄ ⁺ regeneration and assimilation were measured by a ¹⁵N isotope dilution method. Daytime regeneration rates at 0–2 m depth were 0–28% of the calculated assimilation rates at ambient NH ₄ ⁺ concentrations. Regeneration was faster during a dinoflagellate bloom in August than in mixed diatom-dinoflagellate blooms in June and September. Most of the NH ₄ ⁺ appeared to be produced by juvenile copepods, rotifers, tintinnids, and heterotrophic dinoflagellates in the size fraction 45–200 m.     

Seasonal effects of light,temperature, nutrient concentration and salinity on the physiology and growth of Caulerpa paspaloides (Chlorophyceae)

Caulerpa paspaloides (Bory) Greville were collected during the winter and summer (1978 to 1979) from the Florida Keys, USA. Thalli collected during the winter photosynthesized more efficiently at low light intensities (I_c<1, I_k=38 Exm^-2xs^-1) than did thalli collected in the summer (I_c=13, I_k=111 Exm^-2xs^-1). Summer thalli exhibited higher P_max values (2.20 mgO₂xg^-1 dry wtxh^-1) than winter thalli (1.70 mg O₂xg^-1 dry wtxh^-1). Rates of rhizome elongation and frond initiation were strongly inhibited by winter temperatures. The maximum lethal temperature for summer thalli was 37° to 38°C as measured by both growth and photosynthesis. No evidence of nitrogen or phosphorus limitation was found. Relatively minor reductions in salinity (3S) resulted in significant increases in rhizome apex motality. Results indicate that low winter temperatures are responsible for reduced winter growth rates previously reported for the Key Largo population. Increased photosynthetic efficiency at low light intensities and preferential maintenance of rhizome elongation over frond initiation appear to allow this tropical macroalga to optimize growth and survival during the winter.     

Light-saturated photosynthesis by phytoplankton size fractions in the New York bight,USA

The relationships between netplankton and nanoplankton assimilation numbers, temperature, and major nutrient concentrations were studied and evaluated in the context of seasonal patterns in the biomass of these phytoplankton size fractions. Netplankton and nanoplankton blooms typically occur during late winter (2° to 8°C) and summer (18° to 24°C), respectively. Variations in nanoplankton and netplankton assimilation numbers were not statistically related to the development or collapse of specific blooms based on weekly sampling, but assimilation numbers were higher during the bloom periods than during transition periods of rapid temperature change (8° to 18°C). Differences in the assimilation numbers between size fractions could account for the dominance of the nanoplankton fraction during the summer bloom period but not for the dominance of netplankton during the winter bloom period. Nanoplankton and netplankton assimilation numbers were exponential functions of temperature between 8° and 24°C and 8° and 20°C, respectively. Below 8°C the assimilation numbers of both fractions were higher than expected on the basis of temperature. Above 20°C netplankton assimilation numbers declined with temperature. Netplankton and nanoplankton assimilation numbers were occasionally correlated with dissolved inorganic nitrogen concentrations from less than 1.0 to more than 15 g-at l^-1. Under these conditions, nanoplankton growth rates (calculated from assimilation number and carbon:chlorophyll) were higher and increased more rapidly with dissolved inorganic nitrogen than netplankton growth rates.     

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.     

Ecological growth strategies in the seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae): Soluble nitrogen and reserve carbohydrates

The seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae) were grown in an outdoor continuous-flow system at both ambient incident light (I₀) and 0.13 I₀. During the winter, both species accumulated substantial soluble nitrogen reserves (up to 1020 g-at N·g dry wt^-1 in G. foliifera and 630 g-at N·g dry wt^-1 in Ulva sp.). The rate at which these N reserves were depleted was proportional to the growth rate. Seaweeds grown at 0.13 I₀ had lower growth rates and higher levels of soluble tissue N than plants grown at I₀. During the spring-summer growing season, peaks in tissue N followed nutrient peaks in the ambient seawater. Ulva sp. had higher nutrient uptake and growth rates than G. foliifera and showed greater fluctuations in soluble tissue N. This may characterize opportunistic seaweed species with high biomass turnover rates. At I₀, the levels of starch (up to 340 mg·g dry wt^-1 in G. foliifera and 170 mg·g dry wt^-1 in Ulva sp.) were highest during the spring and summer. During this period, fluctuations in starch content were inversely related to growth rate and soluble tissue N. Seaweeds grown at 0.13 I₀ did not accumulate starch. Neither species was found to overwinter with starch reserves.     

Persistent blooms of surf diatoms along the Pacific coast,USA

Productivity was studied in two diatom species, Chaetoceros armatum T. West and Asterionella socialis Lewin and Norris, which form persistent dense blooms in the surf zone along the Pacific coast of Washington and Oregon, USA. Past observations have shown that surf-diatom standing stock usually declines in summer along with concentrations of nitrate and ammonium. Using the ¹⁴C method, photosynthetic rates in natural surf samples were measured monthly for one year (October 1981 through September 1982) at a study site on the Washington coast. Also measured were temperature, salinity, dissolved nutrients, particulate carbon and nitrogen (used as estimates of phytoplankton C and N), and chlorophyll a. Assimilation numbers (P _max) were higher in summer (5 to 8 g C g^-1 chl a h^-1) than in winter (3 to 4gC). Specific carbon incorporation rates (µ_max) showed no obvious seasonality, mostly falling within the range of 0.09 to 0.13 g C g^-1 C(POC) h^-1. The discrepancy between the seasonal trends for chlorophyll-specific and carbon-specific rates reflects a change in the carbon-to-chlorophyll ratio. Because of seasonal differences in daylength and light intensity, daily specific growth rates () are thought to be higher in summer than in winter. Neither ammonium enrichment assays nor particulate carbon-to-nitrogen ratios provided convincing evidence for nitrogen limitation during summer, and the observed changes in diatom abundance cannot be explained on this basis. Both the high diatom concentrations and their seasonal variations probably are due mainly to factors other than growth rates; two factors considered important are diatom flotation and seasonal changes in wind-driven water transport. C. armatum usually dominates the phytoplankton biomass in the surf zone, and evidence suggests that this species is strongly dominant in terms of primary production.Contribution No. 1391 of the School of Oceanography, University of Washington, Seattle, Washington, USA     

Growth of juvenile Macrocystis pyrifera (Laminariales) in relation to environmental factors

Instantaneous relative growth rates, (d^-1), were measured for juveniles of the giant kelp Macrocystis pyrifera transplanted to study sites in Southern California kelp forests between 1978 and 1982. Growth rates ranged from negative values (indicating loss of tissue) to 0.03 (doubling of total frond length every 19 d). Multiple regression analysis of growth versus irradiation, temperature, nitrogen concentration and amount of fouling revealed that all these factors had significant effects, together accounting for about 50% of the total variance. Elevated irradiation and nitrogen levels had strongly stimulatory effects (tissue nitrogen may have been more critical than ambient nitrogen for growth), while high temperature and fouling had strongly inhibitory effects. Irradiation was the most important factor influencing growth in 6 of the 8 transplant experiments. During these 6 experiments, the compensating irradiation level (below which there was no growth) was between 0.4 and 0.7 E m^-2 d^-1, and saturating irradiation was between 2 and 3 E m^-2 d^-1. During two of the experiments, growth was apparently limited by extremely high temperatures or low nitrogen levels. Quantum irradiation levels in the kelp forest were generally between the compensation and saturation levels. However, irradiation levels occasionally dropped below the compensation point for several months. Irradiation was occasionally low enough to limit the distribution of juvenile kelp by inhibiting growth, especially in the deeper portions of the kelp forest and under dense canopies formed by adult plants.     

Application of a 15N tracer method to the study of dissolved organic nitrogen uptake during spring and summer in Chesapeake Bay

The dissolved organic nitrogen (DON) pool in marine waters contains a diverse mixture of compounds. It is therefore difficult to accurately estimate planktonic uptake of DON using the limited number of radiolabeled compounds commercially available. We describe a method to estimate DON uptake rates using ¹⁵N-labeled DON recently released from phytoplankton. To make ¹⁵N-labeled DON, we incubated surface water with ¹⁵NH ₄ ⁺ and then isolated the DON, including any recently released DO¹⁵N, with ion retardation resin. This DON was then added to a freshly collected water sample from the same environment to quantify the rate of DON uptake. The technique was applied to investigate rates of DON uptake relative to inorganic nitrogen in the mesohaline Chesapeake Bay during May 1990 and August 1991. The May experiment took place after the spring bloom, and rates of DON uptake [ranging from 0.31 to 0.53 g-atom (g-at) Nl^-1 h^-1] often exceeded rates of NH ₄ ⁺ and NO ₃ ^- uptake combined. The rates of DON uptake at this time were higher than estimated bacterial productivity and were not correlated with bacterial abundance or bacterial productivity. They were, however, correlated with rates of NO ₃ ^- uptake. In May, we estimate that only 7 to 32% of DON uptake was a result of urea utilization. In contrast, in August, when regenerated nutrients predominate in Chesapeake Bay, rates of DON uptake (ranging from 0.14 to 0.51 g-atom Nl^-1 h^-1) were an average of 50% of the observed rates of NH ₄ ⁺ uptake. Consistent with the May experiment, rates of DON uptake were not correlated with bacterial production. A sizable fraction of DON uptake, however, appeared to be due to urea utilization; rates of urea uptake, measured independently, were equivalent to an average of 74% of the measured rates of DON uptake. These findings suggest that, during both periods of study, at least a fraction of the measured DON uptake may have been due to utilization by phytoplankton.     

Occurrence and significance of nitrogen transport in the brown alga Laminaria digitata

Nitrogen transport in a Laminaria digitata (Huds.) Lamour population growing at Arbroath, Scotland has been investigated (during the period 3 June 1981 to 3 July 1981), using ¹⁵N as tracer. NO ₃ ^- was assimilated both by the blade meristem and by the mature blade. NO ₃ ^- uptake by the blade meristem alone was insufficient to supply the nitrogen demand for growth. This additional demand was met by nitrogen transport from the mature blade to the meristem. It was estimated that 70% of the nitrogen demand of the meristematic region was supplied by nitrogen transport from the mature blade. Although transport occurred, the size of the endogenous pool of stored nitrogen in the mature blade did not change, the quantity of nitrogen transported being equivalent to the amount of NO ₃ ^- assimilated during the experimental period. The possible co-transport of nitrogen and carbon is discussed.     

Carbon and nitrogen dynamics during growth and degradation of phytoplankton under natural surface irradiance

Under conditions of natural irradiance, the development and decline of a flagellate-dominated phytoplankton population was followed in a coastal North Atlantic pond over a 3 d period in summer 1986. Irradiance negatively affected phytoplankton biomass estimated as chlorophyll a, which decreased during the day at photosynthetically available radiation (PAR) levels above 600 to 1000 mol m^-2s^-1; chlorophyll a increased at PAR values below this threshold. In addition, an inverse relationship was found between changes in chlorophyll a and changes in dissolved inorganic nitrogen, indicating synthesis of nitrogenous biomass mainly at night and degradation mainly during the day, with intense exchanges of material between the particulate and dissolved nitrogen fractions. The natural abundance of ¹³C in particulate matter increased initially, and then remained constant, and was controlled mainly by the ratio -carboxylases activity: ribulose biphosphate carboxylase activity. The hypothesis that the latter enzyme is broken down under high irradiance and is partly responsible for increases in external dissolved nitrogen was rejected.     

Growth patterns of Codium fragile ssp. tomentosoides in response to temperature,irradiance, salinity,and nitrogen source

Seasonal patterns of growth, reproduction, and productivity of Codium fragile spp. tomentosoides (van Goor) Silva were monitored at 3 locations in Rhode Island. Maximal growth occurred during the summer and was more significantly correlated with temperature than any other factor measured in this study. Multiple correlation models suggested an interaction between temperature, irradiance, and available nitrogen. Maximal reproduction occurred in late summer and early fall. The maximal productivity, based on harvested quadrats, was 2. 10 g dry weight m^-2 day^-1. A large amount (up to 87.3%) of the annual production entered the detrital food chain during the winter by fragmentation of the thallus. Culture studies examined the effects of temperature (6° to 30°C), irradiance (7 to 140 E m^-2 sec^-1), daylength (8 h light: 16 h dark to 24 h light: O h dark) and salinity (6 to 48) on growth. Differentiated thalli grew over a broad range of experimental conditions, with maximal growth at 24°C, 24 to 30 S, a minimal irradiance of 28 E m^-2 sec^-1, and 16 h daylength. The effect of increasing daylength was due to increased total daily irradiance rather than to a true photoperiodic effect. Undifferentiated sporelings survived and grew in a narrower range of environmental conditions than thalli. c. fragile spp. tomentosoides grew equally well with nitrate, nitrite, ammonium, and urea as a nitrogen source. The addition of NaHCO₃ stimulated growth at levels of 2.4 to 4.8 mM, suggesting an inorganic carbon limitation in static cultures. This study supports the hypothesis that the in situ seasonal growth pattern of c. fragile spp. tomentosoides is primarily due to the interaction of temperature and irradiance.     

Isotopic characterization of atmospheric nitrogen inputs as sources of enhanced primary production in coastal Atlantic Ocean waters

Current estimates indicate that atmospheric nitrogen deposition is responsible for 26 to over 70% of new nitrogen (N) input to North Carolina estuaries and coastal waters. Concentrations of N in coastal rainfall events in a 2-yr period (August 1990 to 1992) ranged from 0.7 to 144 M for NO ₃ ^- and 0.5 to 164 M for NH ₄ ⁺ . The ¹⁵N values of the NO ₃ ^- and the NH ₄ ⁺ were determined in 15 rain events. NH₄ ⁺ values averaged-3.13 (range:-12.5 to+3.6), while NO ₃ ^- plus dissolved organic N fractions had an average ¹⁵N of+1.0 (range:-2.0 to+4.7). The uptake of this isotopically light N into particulate N, in parallel with primary productivity and biomass (as chlorophyll a) determinetions, was examined in microcosm and mesocosm bioassays. As phytoplankton productivity and biomass increased with added rainwater N, the ¹⁵N of particulate N decreased. To investigate the effects of significant atmospheric N loading with stable isotope tracers, we measured the ¹⁵N of the>1 m fraction from surrounding coastal waters. Owing to the episodic nature of atmospheric deposition and the great variation in N loading with each event, a simple assessment of the atmospheric contribution was not possible. During a period in which rainfall inputs were significant and frequent (August 1992), ¹⁵N values were several more negative than during periods of drought (Fall 1990). These experiments and observations emphasize the contribution of atmospheric nitrogen deposition to new production in coastal waters.