Numerical behavior of a zooplankton,phytoplankton and phosphorus system

Several numerical features of a three-dimensional dynamical system which models the three species system made up of phytoplankton, zooplankton and organic phosphorus nutrient in a lake environment are presented. Certain properties of this system, such as the existence of limit cycles, are demonstrated numerically. It is shown that in certain time domains the system may be stiff, thus requiring the use of suitable algorithms.     

Optimization of exergy and implications of body sizes of phytoplankton and zooplankton in an aquatic ecosystem model

Size appears to be an important parameter in ecological processes. All physiological processes vary with body size ranging from small microorganisms to higher mammals. In this model, five state variables — phosphorus, detritus, phytoplankton, zooplankton and fish are considered. We study the implications of body sizes of phytoplankton and zooplankton for total system dynamics by optimizing exergy as a goal function for system performance indicator. The rates of different sub-processes of phytoplankton and zooplankton are calculated, by means of allometric relationships of their body sizes. We run the model with different combinations of body sizes of phytoplankton and zooplankton and observe the overall biomass of phytoplankton, zooplankton and fish. The highest exergy values in different combinations of phytoplankton and zooplankton size indicate the maximum biomass of fish with relative proportions of phytoplankton and zooplankton. We also test the effect of phosphorus input conditions corresponding to oligotrophic, mesotrophic, eutrophic system on its dynamics. The average exergy to be maximized over phytoplankton and zooplankton size was computed when the system reached a steady state. Since this state is often a limit cycle, and the exergy copies this behaviour, we averaged the exergy computed for 365 days (duration of 1 year) in the stable period of the run. In mesotrophic condition, maximum fish biomass with relative proportional ratio of phytoplankton, zooplankton is recorded for phytoplankton size class 3.12 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume). In oligotrophic condition the highest average exergy is obtained in between phytoplankton size 1.48 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume), whereas in eutrophic condition the result shows the highest exergy in the combination of phytoplankton size 5.25 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume).     

Seston composition of the bottom waters of Great Lameshur Bay,St. John,US Virgin Islands

Seston composition [particulate organic carbon (POC), particulate nitrogen (PN), phyto- and microzooplankton numbers and biomass] was investigated in the bottom waters of Great Lameshur Bay, St. John, Virgin Islands (USA), in October, 1970 during Tektite II Mission 17–50. Mean values of 67.4 g POC · I^-1 and 7.2 g PN · I^-1 were determined. A mean phytoplankton carbon content of 42.2 g · I^-1 and zooplankton carbon content of 5.5 g · I^-1 were calculated from counts. The phytoplankton consisted mainly of dinoflagellates 71.2% phytoplankton carbon. Copepods were the dominant zooplankters (61.8% zooplankton carbon), followed by larvaceans (30.9% zooplankton carbon). Organic carbon content of counted zooplankton faecal pellets ranged between 0.4 and 1.6 g · I^-1, and amounted probably to about 15% of the total zooplankton carbon value. Plankton and detritus components as possible food for coral-reef animals are discussed. The ratio carbon: nitrogen of suspended particles is compared to that of sedimented matter.     

Mechanisms of production and fate of organic phosphorus in the northern Adriatic Sea

In the period from 1980 to 1984 organic phosphorus, nutrients, primary production rates (¹⁴C), chlorophyll a (chl a) standing crops, and basic oceanographic parameters were measured during 23 cruises at six stations in the open waters of the northern Adriatic Sea. These waters are significantly influenced by polluted Po River discharge. Organic phosphorus was correlated with several parameters which characterize phytoplankton activity and organic matter decomposition processes. In the late winter-spring period, organic phosphorus is produced during phytoplankton blooms. It is hypothesized that microzooplankton grazing is the main factor increasing the organic phosphorus concentrations in summer (up to 1.1 mol 1^-1). Fall and winter had much lower values (below 0.3 mol 1^-1) due to remineralization processes and an increased water mass exchange between the northern and central Adriatic regions. The direct contribution of organic phosphorus by freshwater discharge was not found to be significant. The higher organic phosphorus concentrations that can occur in low salinity waters are most likely due to their increased capability to support primary production.     

The common jellyfish Aurelia aurita: standing stock,excretion and nutrient regeneration in the Kiel Bight,Western Baltic

The population dynamics, ammonia and inorganic phosphate excretion, and nutrient regeneration of the common jellyfish Aurelia aurita was investigated from 1982 to 1984 in the Kiel Bight, western Baltic Sea. During summer 1982, medusae abundance ranged between 14 and 23 individuals 100 m^-3, biomass was estimated at about 5 g C 100 m^-3 and the mean final diameter of individuals was 22 cm. Abundance, based on numbers, in 1983 and 1984 was an order of magnitude lower; biomass was less than 2 g C 100 m^-3 and jellyfish grew to 30 cm. During the summers of 1983 and 1984, A. aurita biomass constituted roughly 40% of that of the total zooplankton>200 m. In 1982, for which zooplankton data were lacking, it was assumed that medusae biomass was greater than that of all other zooplankton groups. Total ammonia excretion ranged between 6.5 and 36 mol h^-1 individual^-1, whereas inorganic phosphate release was 1.4 to 5.7 mol h^-1 individual^-1. Allometric equations were calculated and exponents of 0.93 for NH₄–N release and 0.87 for PO₄–P excretion were determined. Nitrogen and phosphorus turnover rates were 5.4 and 14.6% d^-1, respectively. In 1982, the medusae population released 1 100 mol NH₄–N m^-2 d^-1, about 11% of the nitrogen requirements of the phytoplankton. The inorganic phosphate excretion (150 mol m^-2 d^-1) sustained 23% of the nutrient demands of the primary producers. In the other two years the nutrient cycling of the medusae was much less important, and satisfied only 3 to 6% of the nutrient demands. It is suggested that in some years A. aurita is the second most important source of regenerated nutrients in Kiel Bight, next to sediment.     

Excretion of ammonia by zooplankton and its potential contribution to nitrogen requirements for primary production in the Catalan Sea (NW Mediterranean)

Excretion of ammonia by mesozooplankton (>200 m zooplankton) and its potential contribution to the nitrogen requirement for phytoplankton growth has been estimated for different hydrographical situations along a transect across the Catalan Sea (Northwestern Mediterranean). The nitrogen excreted as ammonia was estimated from mesozooplankton biomass and specific excretion rates. Nitrogen requirements of phytoplankton were estimated by means of carbon fixation rates and C:N ratios of <200 m particulate organic matter. Minimum C:N ratios and maximum primary production, zooplankton biomass, phytoplankton nitrogen requirements, and nitrogen excretion of zooplankton occurred near the Catalan density front. On average, the nitrogen regenerated by the mesozooplankton accounted for 43% of the nitrogen requirements of the phytoplankton. The specific excretion rates of ammonia and the percentage of phytoplanktonnitrogen requirements supplied by excreted nitrogen were higher at coastal stations. In some coastal and frontal stations, the ammonia excreted exceeded the phytoplanktonnitrogen demand. Bacteria competing for nutrient supply and the possible uncoupling between rate processes and standing stocks of phyto- and zooplankton could explain the apparent excess of regenerated ammonia.     

Zooplankton abundance and grazing at Davies Reef,Great Barrier Reef,Australia

Zooplankton abundance and grazing on autotrophic and heterotrophic particulate matter were measured along a transect across Davis Reef (18°5S; 147°39E) and in the back-reef lagoon over tidal and diel cycles during austral winter (August 1984). Zooplankton entering the reef from the surrounding shelf waters decreased in abundance over the reef flat, presumably because of predation. Within the reef lagoon, maximum daytime densities of pelagic copepods occurred during high water, suggesting an external input. At night, water-column zooplankton biomass increased by a factor of 2 to 3 due to the emergence of demersal reef zooplankton. Zooplankton grazing rates on heterotrophic particulate matter (bacteria + detritus and Protozoa) compared to phytoplankton were higher on the reef flat than on the fore-reef or lagoon. Within the lagoon, zooplankton grazing rates on heterotrophic material were maximum during high water, coincident with maximum tidal concentrations of particulate organic carbon. The combined demersal and pelagic zooplankton community were often able to crop 30% of the daily primary production by >2µm phytoplankton. However, >50% of phytoplankton biomass was in cells <2µm, presumably unavailable to these zooplankton. Our particulate production and ingestion measurements, together with zooplankton carbon demand extrapolated from respiration estimates, suggest that the zooplankton community of Davies Reef derives much of its nutrition from detritus.Joint contribution from the University of Maryland, Center for Environmental and Estuarine Studies (No. 2015), and the Microbial Ecology on a Coral Reef Workshop (MECOR No. 19)     

Analyse des paramètres hydrobiologiques dans la remontée de Cabo Frio (Brésil)

South Atlantic Central Waters (SACW) upwell close to Cabo Frio (Brazil, Rio de Janeiro) shore. The resultant typical hydrobiological conditions were studied at an anchorage station over an annual cycle from February 1973–February 1974. Multivariate analyses of hydrological and planktonic data revealed the structure of the ecosystem and the factorial relations governing its dynamics. Alternation, superposition or mixing of the water masses of different origin (Brazil current, coastal, SACW) constitute the most important factors responsible for the great hydrobiological variability observed in the study area. Spasmodic changes in wind direction and force are superimposed on a seasonal trend of more frequent upwelling in summer than during the rest of the year. The deep water is characterized by temperatures of <18°C, nitrate contents of 10 g-at l^-1 and by organic matter mainly composed of detritic elements from the shelf. Temperature variations together with nutrient contents (NO₃ or PO₄) reflect variations in primary biomass at the surface but not at 50 m depth, where detrital matter precluded valid measurements. Water of the Brazilian Current (salinity 36.0) frequently mixes with deep water of the thermal front, or with coastal water (<35.0) which invades the area when south-west winds prevail. This lower-salinity water is rich in seston particles. During the study period, primary biomass was relatively low due to eutrophication. We observed less than 3 g l^-1 chlorophyll and 10⁶ phytoplankton cells per litre: the phytoplankton populations were highly diversified, indicating an advanced degree of complexity and evolution of this ecosystem. The observed formation of a thermocline constitutes an important inducing factor for an algal bloom. Simultaneous phyto- and zooplankton maxima would induce an increased grazing rate by herbivorous zooplankton which would also partly explain the relatively low level of primary biomass. Zooplankton is as abundant here as in other great upwelling regions: 100 organisms l^-1 and 200 mg organic matter m^-3.     

Zooplankton of tikehau atoll (Tuamotu archipelago) and its relationship to particulate matter

The standing stock and taxonomic composition of zooplankton (>200 m) were monitored in the lagoon of Tikehau atoll from April 1985 to April 1986. These data were supplemented by two 10 d studies on the variability, structure and functioning of the pelagic ecosystem. The biomass of animals >200 m comprised 50% of the total biomass of all organisms from 35 to >2000 m. The zooplankton populations were characterized by successive blooms of copepods, larvaceans, pteropods and salps, probably arising from the periodic input of detritus from the reef during windy periods. As a result, the ecosystem was not in a steady state and the data for the fluxes of organic matter are presented separately for April 1985 and April 1986. Using the C:N:P ratio method, net growth efficiencies, K ₂, were calculated for total mesozooplankton, mixed copepods, and two planktonic species, Undinula vulgaris and Thalia democratica. Combined with nitrogen and phosphorus excretion rates, these K ₂ values enabled the assessment of production rates. On a 24 h basis, P:B ratios (%) were close to 100 for the total zooplankton and 54, 34 and 800 for mixed copepods, U. vulgaris and T. democratica, respectively. These ratios are 5.7 times lower than that recorded for phytoplankton. High productivity may be ascribable to the high density of seston, the high temperature (29.5°C), and the kind of organisms present. Zooplankton production equalled 38 and 30% of ¹⁴C uptake during April 1985 and April 1986, respectively. Ingestion of animals >35m was calculated by means of assimilation efficiencies and amounted to 17 and 7% of particulate organic carbon, 100 and 38% of living carbon, and 64 and 140% of primary production during the two periods, respectively. Finally, inorganic exduring was 32 and 18% of phytoplankton nitrogen and phosphorus requirements. A model based on the dimensional structure of the pelagic food-web, has been drawn to illustrate the biomass and carbon, nitrogen and phosphorus fluxes in the study area. The lagoon appears to export part of its planktonic biomass, which is 4.2 times lower one sea mile outside the main pass connecting the lagoon to the open ocean.     

Phosphorus dynamics associated with phytoplankton blooms in eutrophic Mikawa Bay,Japan

Changes in the phosphorus components of the particulate matter in seawater were studied in the eutrophicated waters of Mikawa Bay, Japan, during summer 1981. The contents of particulate phosphorus and hot-water extractable intracellular phosphorus displayed remarkable changes associated with phytoplankton blooms caused by wind-induced or upwelling-associated nutrient enrichment from the bottom water layers. Nanoplankton <10 m accounted for much of the particulate phosphorus (70 to 79% in June and July, and 44 to 78% in August and September); the contribution of large-sized phytoplankton >25 m varied from 9 to 49%, the peak values being attained under red-tide conditions. The capacity for phosphorus storage in cells was low in nanoplankton cells, high in large phytoplankton species. Differences in rates of phosphorus storage and growth between nanoplankton and large phytoplankton accounted for fluctuations in particulate phosphorus which were closely associated with fluctuations in phytoplankton blooms in Mikaw Bay.     

Sedimentation of particulate matter during a phytoplankton spring bloom in relation to the hydrographical regime

Data presented and discussed here were collected continuously during April/May 1975 in the Bornholm Basin of the Baltic Sea. Sedimentation rates of particulate matter were recorded with 5 multisample sediment traps from different depths in the water column at 2 positions 170 km apart. Current meter data collected during the same period and depths indicated that the positions remained hydrographically distinct during the investigation. Particulate matter from the euphotic zone including diatom cells formed the bulk of the material collected by all traps. This flux of organic particles to the bottom was unimpeded by the strong density stratification present in the water column. The upper traps always collected less material than lower ones. This paradox has been ascribed to diminishing current speeds with depth, concomitant with an increase in sinking rates of phytoplankton and phytodetritus. Both factors influence the sampling efficiency of sediment traps, which are thought to have underestimated actual sedimentation rates here. A time lag of 2 to 3 weeks in bloom development seemed responsible for the characteristic differences between the two positions. The phase of major sedimentation at one position covered about 18 days, and a distinct sequence in the composition of the material collected by the 6 glasses of each trap indicated phases of a progressively deteriorating phytoplankton population in the water column contributing the particulate material. A total of 6.2 g C m^-2 in 34 days was recorded at this station. Apart from a trap situated in an oxygen deficient layer which collected 0.44 g C m^-2 of zooplankton corpses, zooplankton mortality was overestimated by the traps. Large-scale sedimencation of fresh organic matter produced by the spring bloom is probably a regular feature in areas with low over-wintering zooplankton populations and, as such, possibly has a direct stimulatory effect on growth and reproduction of the benthos.Contribution No. 185 of the Joint Research Programme 95, Kiel University.     

A vertical-compressed three-dimensional ecological model in Lake Taihu, China

A three-dimensional ecological model on the basis of the analyses of environmental characteristics is set up for Lake Taihu, one of the largest shallow lakes in China. The hydrodynamic processes, nutrient cycling, chemical processes and biological processes are integrated in the model. Model state variables include: water current, surface displacement, nutrients of nitrogen and phosphorus, as well as their different forms such as ammonia nitrogen, nitrate nitrogen, phosphate phosphorus, etc., biomasses of macroplankton, phytoplankton, zooplankton and fish, and also the nutrient levels of macroplankton and phytoplankton. A nutrient budget and sediment transformation are also coupled in the model. The data from January 17, 1997 to January 18, 1998 are use to calibrate the model. The model results have shown good agreement with the observations. It implies that the model could be used for the lake environmental management and research for examining the processes and determining the water quality. The reasons of deviations between the modelled results and the observed values are also discussed. There are six factors that explain the deviations of the modelled results from the observed values and they can be grouped into two sets. One set of problems is associated with the standard deviation introduced by sampling and analyses. The second set of problems can be solved by introduction of processes lacking in the present model (resuspension, phytoplankton transportation mode under the wind with low speed, shifts in species composition and varied size of phytoplankton and zooplankton). The latter two processes should be included in the model at a later stage by integration of a structurally dynamic approach into the three-dimensional model.     

Phytoplankton carbon dynamics during a winter-spring diatom bloom in an enclosed marine ecosystem: primary production,biomass and loss rates

Phytoplankton production, standing crop, and loss processes (respiration, sedimentation, grazing by zooplankton, and excretion) were measured on a daily basis during the growth, dormancy and decline of a winter-spring diatom bloom in a large-scale (13 m³) marine mesocosm in 1987. Carbonspecific rates of production and biomass change were highly correlated whereas production and loss rates were unrelated over the experimental period when the significant changes in algal biomass characteristic of phytoplankton blooms were occurring. The observed decline in diatom growth rates was caused by nutrient limitation. Daily phytoplankton production rates calculated from the phytoplankton continuity equation were in excellent agreement with rates independently determined using standard ¹⁴C techniques. A carbon budget for the winter bloom indicated that 82.4% of the net daytime primary production was accounted for by measured loss processes, 1.3% was present as standing crop at the end of the experiment, and 16.3% was unexplained. Losses via sedimentation (44.8%) and nighttime phytoplankton respiration (24.1%) predominated, while losses due to zooplankton grazing (10.7%) and nighttime phytoplankton excretion (2.8%) were of lesser importance. A model simulating daily phytoplankton biomass was developed to demonstrate the relative importance of the individual loss processes.     

Estimating carbon flux to pelagic grazers in the ice-edge zone of the eastern Bering Sea

Zooplankton ingestion of phytoplankton carbon in the iceedge zone of the Eastern Bering Sea was measured using a deck incubation approach in 1982. Using further samples collected in 1983, the plant cell carbon to cell volume ratio was estimated at 0.0604 pg m^–3 from an experimentally determined particulate carbon to seston volume relationship. The application of this conversion to the results of experimental incubations of natural plant stocks with net-caught zooplankton produced ingestion rates of 68.8 and 10.26 mg C g^–1 grazer d^–1 for copepods and euphausiids, respectively. Extrapolating these rates to in situ zooplankton biomass at the edge of the seasonal ice pack yielded carbon flux rates through the zooplankton community ranging between 6.5 and 32.8 mg C m^–2 d^–1. This consumption amounted to less than 2% of the daily phytoplankton production in the ice-edge zone.     

Marine food-web analysis: An experimental study of demersal zooplankton using isotopically labelled prey species

Short-term incubations in seawater containing H¹⁴CO₃ ^- or ³H₂O in place of the naturally predominant isotopes can yield highly radioactive preparations of living phytoplankton or zooplankton. Subsequent in situ incubation of these labelled organisms with the community from which they were taken results in the rapid transfer of radioisotope to those species which prey upon them. This technique has been employed to map a portion of a marine food web involving demersal zooplankton; experiments were conducted in summer and autumn on a coral reef and in a subtropical estuary. Similar results were obtained from these initial experiments at each study site during both seasons. Prey supplied as zooplankton (124 to 410 m nominal diameter), which consisted mainly of Oithona oculata, was fed upon by zooplankton size classes ranging from 410 to 850 m and containing amphipods, ostracods, cumaceans and polychaetes. In experiments employing labelled phytoplankton as prey a wide size spectrum was used (10 to 106 m) in order to include representative samples of most of the available planktonic autotrophs as estimated by primary production measurements. In two separate experiments, only 7 out of 63 samples evidenced grazing of phytoplankton by demersal zooplankters. In contrast, labelled diatom auxospores, employed in one experiment as they constituted the most numerically abundant species in the water column, were found to be grazed upon in nearly half the samples examined.     

On plankton production in Kungsbacka Fjord,an estuary on the Swedish west coast

Environmental conditions, primary production, and zooplankton populations were studied from May, 1969 to November, 1970 at one station in Kungsbacka Fjord, Sweden. The fjord, with an arca of 53 km², is a moderately polluted estuary, with a small tidal range. Data for primary production and environmental parameters were correlated using Spearman's rank correlation coefficient. The annual rate of primary production in 1970 was about 100 gC·m^-2. Carbon fixation was about 80 g·m^-2 in May–November in1969 and 1970. The average monthly rate was highest in June, 1970, with 25 gC·m^-2; about 15 gC·m^-2 was recorded in August–October of both years. Carbon fixation by the phytoplankton was estimated to be about 2,800 tons in the whole fjord in 1970. The average fresh-water inflow to the fjord, amounting to about 13 m³·sec^-1, added about 380 tons of organic carbon, 45 tons of nitrogen, and 4.5 tons of phosphorus per month. Primary production displayed strong correlation with temperature at different depths (P<0.05 to 0.001), indicating the sediments to be the most important nutrient source. A total of 19 holoplanktonic zooplankton species was identified, copepods being the dominant group. The highest zooplankton biomass, 800 to 900 mg·m^-3, was recorded in June of both years. The production of copepods in May–October was about 1 gC·m^-2 in both years. The total secondary production of the zooplankton was calculated as only 1.8 gC·m^-2 in 1970.     

Significance of ammonia in determining the N:P ratio of the sea water off Barbados,West Indies

N:P atomic ratios calculated on NO₃-N alone for the upper waters of the tropical Atlantic Ocean off Barbados are very low, being only 9.8:1. Absolute values are also low, the integrated values between O and 100 m for NO₃-N and PO₄-P being 0.59 and 0.06 g-at l^-1, respectively. However, when ammonia is included as a nitrogen source the ratio becomes 28.8:1. This is the average value obtained from 42 samples taken over a 21-month period, and suggests that phosphorus, and not nitrogen, is the more critical nutrient in phytoplankton growth off Barbados.     

Production organique primaire dans un milieu saumâtre eutrophe (Etang de Berre). Effets d'une forte dilution (dérivation des eaux de la Durance)

¹⁴C primary production measurements were made over a period of 5 years (1965–1969, inclusive) in the brackish lake Etang de Berre, near Marseilles (France). The diversion of the River Durance into the Etang de Berre took place during this period (March 1966) and introduced an important modification into the organic production ecosystem, mainly through increased and variable freshening, accompanied by substantial nutrient input. The seasonal distribution of production rates displayed 3 bloom periods: the first (short and slight) in spring, the second (the most important as regards intensity and duration) in summer, and the third in autumn (October). Before the diversion of the river in 1965, the carbon-uptake rates in the lake ranged between 25 mg/m²/day in winter and 800 mg/m²/day in summer-autumn, the mean value for the year being 150 g C/m², which represents 2.5×10⁴ tons of photosynthesized carbon for the whole lake. After the diversion, more than 3000 mg C/m² day were measured; for 1968, the inclusive uptake rate was 384 g C/m², representing 6×10⁴ tons of synthesized carbon for the whole lake. Nevertheless, noticeable variations occurred from one year to another. From the annual nutrient input of phosphate to the Etang de Berre through the inflow of Durance waters, the quantity of potentially synthesizable elements has been calculated, in terms of carbon, according to the normal P:C ratio of organic substances; this quantity is called R. The difference between measured production, P, and R gives a measure of the regenerated production. This portion of production represented about 80% of the total production before 1968 but only 16% in 1969, a year of maximum fresh-water inflow. This phenomenon could be due to modifications of the ecophysiology of the phytoplankton resulting from the considerable freshening. With increasing nutrient load, eutrophication first occurs, then still greater dilution results in inhibition of production.