Summer nutrient dynamics of the Middle Atlantic Bight: nitrogen uptake and regeneration

Nitrate and ammonium uptake and ammonium regeneration rates(by zooplankton, microplankton and benthos) were measured onthe Atlantic continental shelf (Middle Atlantic Bight) duringsummer, 1980. Euphotic zone profiles of NO₃^– and NH₄⁺uptake rates were similar in magnitude and vertical structureover a large geographical area. Microplankton NH₄⁺ regenerationrates, although measured less frequently, also showed a relativelyconsistent vertical structure; rates were positively correlatedwith uptake rates. Nitrate assimilation (‘new’ production)was used to estimate vertical eddy diffusivity and paniculatesinking rates. Eddy diffusion estimates ranged from <0.1to >2.0 cm² s^–1 and were positively related to arealprimary production. Estimated particulate sinking rates averaged5 mg at Nm^–2d^–1 and compared favorably with sedimentationrates measured from free-floating and moored sediment traps.Benthic nitrogen regeneration rates represented <10% of thispaniculate nitrogen flux. Within the mixed layer, NH₄⁺ assimilation(‘regenerated’ production) represented 50–80%of the total (NO₃^– + NH₄⁺ ) nitrogen productivity and33% for the euphotic zone. Of this, 30% was attributed to zooplankton,63% to microplankton (<100 µm) and 7% to benthos. Onthe average, 74% of the microplankton NH₄⁺ regeneration wasassociated with organisms passing 1 µm filters.     

Distribution and ecology of Middle Atlantic Bight Oligochaeta

Distribution and ecology of continental shelf oligochaetes are poorly understood. From 1975 through 1977 baseline ecological studies were conducted in the Middle Atlantic Bight prior to oil exploration activities. A total of 38 species in three oligochaete families were collected. Of this total one family and 29 of the species were undescribed at the time of collection. The most abundant and widely distributed species were Limnodriloides monothecus, L. medioporus, Bathydrilus longus, Phallodrilus biprostatus, and Tubificoides diazi. No single species dominated the oligochaete fauna. All the oligochaete species responded to changes in physical parameters over the shelf similarly to the total macrofauna. Seasonal trends within the oligochaetes were not found. Species that preferred coarser sand sediments were all small interstitial forms. The finer sand and eurytopic species were all burrowers. Zoogeographically the Middle Atlantic Bight oligochaetes were predominantly temperate (16 species) with some southern species (8). Seven species were broadly distributed from the Caribbean to Massachusetts or around the world. Seven species were only known from the area studied.Contribution No. 1397 of the Virginia Institute of Marine Science.     

The closure of the grevelingen estuary: Its influence on phytoplankton primary production and nutrient content

The Grevelingen estuary was cut off from the sea in May 1971, and changed into stagnat lake Grevelingen. After the closure nitrate concentrations decreased to extremely low values (less than 2 μgat NH₃-N/1). Ammonia concentrations varied between 10–30 μgat NH₃-N/1, comparable with the situation before the closure. Phosphate concentrations fluctuated between1–2 μgat PO₄-P/1 in the estuarine phase, and increased to μgat after the closure, presumably caused by decomposition of biological material and release of phosphates from the bottom. Phytoplankton primary production was not markedly affected by the damming up, and amounted to 175 g C/m² in 1971. Communication no. 147 of the Delta Institute for Hydrobiological Research, Yerseke, The Netherlands     

The multiple spawning pattern of weakfish in the Chesapeake Bay and Middle Atlantic Bight

Weakfish Cynoscion regalis were collected from commercial fisheries in the Chesapeake Bay and the Middle Atlantic Bight (n=4380) during 1989–1992 and their reproductive biology assessed using the gonadosomatic index, macroscopic gonad stages, oocyte diameter distributions, microscopic whole oocyte analysis and histology. Sex ratios were approximately 3:1, females to males, in 1990–1992. Most fish (90%) attained sexual maturity by age 1 and at a small size. Estimated mean length at first maturity was: 164mm total length (TL) for males, and 170 mm TL for females. Weakfish spawn within the Chesapeake Bay, as far north as the Virginia/ Maryland border. Although spawning occurred during May–August and gonad development and initiation of spawning was synchronous, cessation of spawning was asynchronous. There was no indication that older fish exhibited a more extended spawning season than younger fish. Weakfish are multiple spawners with indeterminate fecundity. Oocyte development is asynchronous with oocytes of all stages being present in developed ovaries. Because of the complex and dynamic weakfish ovarian cycle, typical methods of assessing reproduction, such as the GSI and macroscopic gonad stages, are inadequate for this species if not used in conjunction with more detailed methods such as histology.     

Seasonal occurrence and dominance of Centropages congeners in the Middle Atlantic Bight,USA

The distribution and abundance of 15 taxa of birds seen from a vessel in the region of Prydz Bay, Antarctica between November and December 1982 are presented. Antarctic petrels were by far the most frequent in the survey area while Adélie and emperor penguins made up 77% of the observed avian biomass. The distribution and abundance of birds was found to be related to ice conditions, air temperature and wind speed but not to water temperature or air pressure. The extent of species associations at sea are shown and the possibilities of diurnal fluctuations in bird numbers at sea are examined.     

Description of phytoplankton and nutrient in spring in the western North Atlantic Ocean

Abundance of diatoms in the northern Sargasso Sea in springhas been observed in a 4-year period between 1981 and 1985 andthen again in 1987. A theory is presented that if stratificationblocks get-through of deep nutrients to the surface, as in thesouthern Sargasso, then in-situ cycling via non-diatoms andammonia dominates. The theory is that if in-situ cycling doesnot dominate, as northward, then stratification does not blockget-through of deep nutrients to the surface, which, in combinationwith the marked growth capacity of diatoms, produce an abundanceof cells. This abundance consumes nitrate, phosphate and silicateat the surface and ultimately this nutrient-depleted water arrivesin the southern Sargasso via the gyre circulation. Thus thenutrient-depleted water of the western Atlantic gyre is consideredto be primarily achieved only by diatoms, in spring and in thenorthern Sargasso. A transect during April 13–16, 1985,north from the Virgin Islands along 63°–64°W showsa great change in the diatom species plus Emiliania huxleyifrom negligible concentrations in the southern half of the sectionto large concentrations in the northern half. This unevennessof distribution can be rephrased by indicating that these speciesare narrow-niched with regard to nutrient availability. Theother species, coccolithophores and dinoflagellates, are evenlydistributed at low concentrations south-north (though some diminishslightly) and so they are broad-niched with respect to nutrient.High species diversity depends on broadness of nutrient nicheand equilibrium conditions of hydrographically stable water(i), implying that when non-equilibrium conditions occur (ii),when niche breadth is less (iii) or both (iv), then diversityis less marked, (i) is confirmed by broad-niched species southward,(ii) is confirmed by broad-niched species northward, (iii) isconfirmed by narrow-niched species southward and (iv) by narrow-nichedspecies northward.     

Spring bloom dynamics in Kongsfjorden,Svalbard: nutrients,phytoplankton, protozoans and primary production

The marine ecosystem in Kongsfjorden (79°N), a glacial fjord in Svalbard, is to a large extent well known with regard to hydrography, mesozooplankton and higher trophic levels. Research on primary production and lower trophic levels is still scare and especially investigations from winter and spring periods. The spring bloom dynamics in Kongsfjorden were investigated in 2002. The development in nutrient conditions, phytoplankton, protozoans and primary production were followed from 15 April until 22 May. The winter/spring in 2002 was categorized as a cold year with sea ice cover and water masses dominated by local winter-cooled water. The spring bloom started around 18 April and lasted until the middle of May. The bloom probably peaked in late April, but break-up of sea ice made it impossible to sample frequently in this period. Diatoms dominated the phytoplankton assemblage. We estimated the total primary production during the spring bloom in 2002 to range 27–35 g C m⁻². There was a mismatch situation between the mesozooplankton and the phytoplankton spring bloom in 2002.     

On primary production in the South Bight of the North Sea

The area of the North Sea next to the Belgian and Dutch coasts has been surveyed since January 1971. The photosynthetic capacity (potential production) of samples collected at 63 stations and 4 depths has been measured using the ¹⁴C technique.

The homogeneity of the water column was demonstrated almost everywhere. A general pattern of decrease of the potential production from the coast to the open sea was demonstrated in all seasons. Figures ranged from 3 to 30 mg C m^-3 h^-1. Primary production was also determined both by calculations, using the Steemann Nielsen (1952) formula, and by in situ measurements. The higher turbidity near the coast has proved to limit considerably the primary production. Figures usually range from 100 to 1500 mg C m^-2 day^-1.

The following topics have also been discussed: relationship between photosynthetic capacity and pigment content, transparency of the water, solar radiation, comparison between production calculated from photosynthetic capacity and production measured in situ, and nutrient uptake.     

Linking population dynamics models with empirically derived models through phytoplankton primary production

There are alternative methods for estimation of phytoplankton primary production (PP) that are fundamentally different in the calculation approach. The process‐oriented PP model is a mechanistic, empirically derived method based on the photosynthesis–light relationships. The population dynamics‐based PP calculation, which is a synthetic method, provides a production estimate based on population dynamics of phytoplankton. These alternative methods were here compared with regard to production estimates and linked to enhance the performance of the existing models of population dynamics applied to a wide variety of lakes worldwide in terms of morphometry, nutrient status, and light environments. Estimates of PP were shown to be sensitive to changes in phytoplankton sinking and zooplankton grazing rates in both methods. Production estimates in the process‐oriented PP model were also sensitive to light‐associated parameters such as day length. Although the production estimated from the population dynamics‐based PP calculation tended to be lower than that from the process‐oriented PP model irrespective of lake morphometry, production estimates calculated from both methods with standard parameterization were comparable when production was estimated on an annual timescale. However, it was also shown that the alternative methods could produce different production estimates when estimated on shorter timescales such as cyanobacterial blooms in summer. Cyanobacteria with low mortality due to grazing and sinking losses have been considered as trophic bottlenecks, but there is increasing evidence that their mortality is, to a considerable extent, due to parasitic pathogens. In the case of cyanobacterial blooms, an addition of parasite‐related loss term (19%–33% of standing stock) resulted in a resolution of the difference in production estimates between the methods. These analyses theoretically support the critical role of parasitism and resolve the bottleneck problem in aquatic ecosystem metabolism.     

Size-fractionated productivity and nutrient dynamics of phytoplankton in subtropical coastal environments

It is now well established that the size distribution of phytoplankton plays an important role in primary production processes and nutrient dynamics of coastal environment. In situ observations showed that nanophytoplankton (3–20 μm) contributed 72.08% and58.18% of phytoplankton biomass and 58.32% and 41.14% of primary productivity to Xiamen Western Waters and the northern Taiwan Strait, respectively; picophytoplankton (0.2–3 μm) dominated the biomass (64.70%) and productivity (66.09%) in the southern Taiwan Strait. Furthermore, nanophytoplankton accounted for 75% of phosphate uptake with the highest rate constant (8.3×10^-5 s^-1) and uptake rate in unit water volume (5.4×10^-5 mmol dm^-3s^-1); picophytoplankton had the highest uptake rate in unit biomass (5.4×10^-5 mmol mg^-1s^-1) and photosynthetic index (3.8 mgC mgChl a^-1h^-1). All the results highlighted the remarkable characteristics of small size ranged (0.2–20 μm) phytoplankton in subtropical coastal environments: main contributor to phytoplankton biomass and production, high efficiency on organic carbon production and nutrient recycling. The far reaching environmental and ecological implications were discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Photophysiology and primary production of phytoplankton in freshwater

In the last 10 years enormous progress has been made in developing new instruments to collect physiological data from natural phytoplankton. In this review we summarize the motivation which has powered this engagement and focus on new technologies used to measure fluorescence, absorption and biochemical compositions of natural phytoplankton cells from nature. Combining the knowledge of phytoplankton structure along with taxon-specific measurements of photosynthetic activity and biochemical cell composition, can lead to new models which increase the reliability of water quality prediction. Furthermore, recent progress in the analysis of photophysiological fitness of phytoplankton cells has revealed new knowledge about the phylogenetic diversity of metabolic strategies to cope with light and nutrient stress. Future progress in single cell analysing systems will be discussed.     

Oxygen Consumption and Dissolved Inorganic Nutrient Production in Marine Coastal and Shelf Sediments of the Middle Atlantic Bight

The benthic oxygen demand and inorganic nutrient release have been investigated in the coastal zone of the Mid-Atlantic Bight of the United States using in situ and ship-board incubations to estimate fluxes across the sediment-water interface. Oxygen demand ranged from ca. 70 ml m⁻² hr⁻¹ nearshore in warm (25°C) muddy sediments to ca. 4 ml m⁻² hr⁻¹ in cold (6°C), coarse-grained sediments offshore. Ammonium ion flux was out of the sediments in all but 4 of 29 incubations, while nitrate was often found to be utilized within or on the sediments, decreasing in 10 of 29 incubations. Those locations with fluxes of nitrate out of the sediment had concentrations in the pore water that were generally high, sometimes up to several mM l⁻¹. Nitrate fluxes were always well below those of ammonia, even where high almost equivalent concentrations of both NH₄ and NO₃ were found at the same locale. On George's Bank only 13 % of total N demand by the phytoplankton was estimated to be supplied by the benthos, whereas in New York Bight the supply amounted to 42 % of the demand. This difference can be attributed to higher pelagic biological and advective (mixing and upwelling) inputs to George's Bank compared to the relatively well stratified New York Bight in late summer and autumn.     

The primary production of phytoplankton in Lake Vechten

The primary production of the phytoplankton of Lake Vechten (The Netherlands) (area, 4.7 ha; mean depth, 6 m), an unpolluted and stratified sandpit was investigated from 1969 to 1980 (except in 1971, 1975 and 1976) by the in situ ¹⁴C-technique. Other data collected include: solar radiation, transparency, oxygen and thermal structure. In winter and spring diatoms, Cryptophyceae and Chlorococcales were important algal groups, while in summer Dinophyceae and Chrysophyceae were important. The chlorophyll-a concentration was compared to the cellular biovolumes (= fresh weight) of the most abundant phytoplankton species. The primary production maxima occurred in winter, spring and during the summer stratification. The vertical profiles of photosynthesis exhibit light inhibition at surface to a maximum of 4 m. The maximum of zooplankton grazing in May–June caused a sharp decrease in the phytoplankton biomass and seston concentration accompanied by the highest transparency (clear water phase).The values for cellular C-fixation range from 10 to 1307 mg C · M^–2 · day^–1 (annual mean of 280 mg C · m^–2 · day^–1). High dark fixation (up to 100%) was encountered in the metalimnion and hypolimnion from July to October together with peaks of ¹⁴C-fixation due to a crowding of phytoplankton and phototrophic anoxic bacteria. Extracellular excretion by phytoplankton, investigated in 1977 to 1979, was 15% of the annual mean of the total C-fixation. The photosynthetic efficiency, turnover rates, and activity coefficients were low, particularly in the summer months when Ceratium hirundinella was predominant. The seasonal variations were controlled mainly by solar radiation and probably phosphate, the former being more important in the non-stratification period and the latter during the stratification period.     

Organic carbon release by phytoplankton: its composition and utilization by bacterioplankton

The present study characterized the rate of production of extracellularlyreleased organic carbon (ROC) by phytoplankton, its molecularweight distribution, subsequent utilization and transformationby bacterioplankton in situ. Primary production rate of phytoplanktonwas high during the study due to continuous blooms of smalldinoflagellates and ranged from 59.8 to 298.7µg CI^–1h^–1. The rate of organic carbon release varied from 1.3to 123.7 µg CI^–1 h^–1 and constituted from4.0 to 68.9% of the total carbon fixed in photosynthesis. TheROC was fractionated on molecular weight (MW) basis. A low MWfraction less than 500 daltons (18.5% of ROC), a fraction ofMW 10 000–30 000 daltons (30% of ROC), and high MW fractionof > 300 000 daltons (15.4% of ROC), were the most dominantin ROC. Bacterioplankton utilized a significant portion of ROC,ranging from 18 to 77%. Part of the utilized ROC incorporatedby the bacterioplankton (31 – 56%), and the remainderwas respired (mineralized). ROC not utilized by bacteria wascomposed of high MW compounds. The dynamics of the in situ utilizationof ROC and its role as a link between autotrophic and heterotrophicprocesses in the estuary are described. ¹Present address: Department of Environmental Microbiology,Institute of Microbiology, Warsaw University, Warsaw, Poland     

A mathematical model of the nutrient dynamics of phytoplankton in a nitrate-limited environment

Insofar as saturation kinetics are applicable to the growth of phytoplankton in laboratory experiments and to growth in nature, the computer modeling of intracellular nutrient partitioning in populations of cells can lead to better understanding of the dynamics of natural populations. A three-compartment mathematical model was developed to represent a phytoplankton population having the capability to store nitrogen in a nitrate-limited environment. Parameters were estimated by fitting the model to data from two chemostat experiments reported by Caperon (1968). The model was used to simulate growth dynamics observed in chemostat and batch experiments. The model demonstrated the changes which may occur in the nitrogenous constituents of a phytoplankton population with time and environmental conditions. The model also demonstrates three phenomena which have been observed in field and laboratory experiments but which are not represented by the customary Monod model: (1) uptake rates may significantly exceed not growth rates, (2) high growth rates may be encountered at very low environmental nitrate concentrations, and (3) the ratio of internal nitrogen to population size may change significantly during a study period. It is suggested that the amount of nitorgen in storage may be used as an indicator of the physiological state of a monospecific population. Parameters for the one-compartment Monod model were estimated by customary methods form data generated by the three-compartment model. It was shown that difficulties encountered in estimating the yield coefficient and the decay coefficient may be attributed to the intracellular storage phenomenon. It was also demonstrated that the one-compartment Monod model was inadequate to accurately represent population growth in chemostat experiments when intracellular storage is a significant factor.     

The carbon and chlorophyll content of phytoplankton from various nutrient regimes

We have measured the relationship between chlorophyll and carbonin phytoplankton cultures and natural populations of phytoplanktonusing simple standard extraction procedures without correctionfor degradation products. During exponential growth of fourcultures of unicellular and colonial green algae and one diatom,chlorophyll varied from 1.22 to 6.08% of the phytoplankton carboncontent (=3.03%). The chlorophyll content of the cultures was lower during nitrogen and phosphorus deficiency.Results from natural populations of phytoplankton from eutrophiclakes and from a coastal marine station gave values mostly rangingfrom 1.5 to 3.7% chlorophyll, corresponding to carbon/chlorophyllratios of 27–67. If only a rough estimate of the phytoplanktoncarbon biomass is required, a simple, efficient extraction procedurecan be used without any corrections for degradation products.     

Nutrient limitation of the primary production of phytoplankton in Lake Baikal

Nutrient limitation of the primary production of phytoplankton at some stations in southern and central Lake Baikal was studied by nutrient enrichment experiments in August 2002. Chlorophyll (Chl.) a concentrations ranged from 0.7 to 5.8μgl⁻¹. Inorganic nutrient concentrations were low: soluble reactive phosphorus ranged from 0.05 to 0.20μmoll⁻¹, ammonia from 0.21 to 0.41μmoll⁻¹, and nitrite plus nitrate from 0.33 to 0.37μmoll⁻¹. In the five enrichment experiments, phosphate spikes and phosphate plus nitrate spikes always stimulated primary production. Nitrate spikes also stimulated primary production in four of the experiments. Significant differences were detected between the controls and phosphate spikes and between the controls and phosphate plus nitrate spikes. Thus, the first limiting nutrient is thought to be phosphorus, but once phosphorus is supplied to the surface water, the limiting nutrient will quickly shift from phosphorus to nitrogen.     

The influence of mesozooplankton on phytoplankton nutrient limitation: a mesocosm study with northeast Atlantic plankton

We used marine phytoplankton from mesocosms seeded with different zooplankton densities to study the impact of mesozooplankton on phytoplankton nutrient limitation. After 7 d of grazing (copepod mesocosms) or 9 d (appendicularian mesocosms) phytoplankton nutrient limitation was studied by enrichment bioassays. After removal of mesozooplankton, bioassay bottles received either no nutrients, phosphorus or nitrogen alone, or a combination of nitrogen and phosphorus and were incubated for 2 d. Phytoplankton reproductive rates in the bottles without nutrient addition were calculated after correction for grazing by ciliates and indicated increasing nitrogen limitation with increasing copepod abundance. No nutrient limitation was found in the appendicularian mesocosms. The increase of nutrient limitation with increasing copepod density seems to be mainly the result of a trophic cascade effect: Copepods released nanoplankton from ciliate grazing pressure, and thereby enhanced nitrogen exhaustion by nanophytoplankton and reduced nitrogen excretion by ciliates. Nitrogen sequestration in copepod biomass, the mechanism predicted by the ecological stoichiometry theory, seems to have been a weaker effect because there was only little copepod growth during the experiment.