Spectral analysis unmasks synchronous and compensatory dynamics in plankton communities

Community biomass is often less variable than the biomasses of populations within the community, yet attempts to implicate compensatory dynamics between populations as a cause of this relationship often fail. In part, this may be due to the lack of appropriate metrics for variability, but there is also great potential for large-scale processes such as seasonality or longer-term environmental change to obscure important dynamics at other temporal scales. In this study, we apply a scale-resolving method to long-term plankton data, to identify the specific temporal scales at which community-level variability is influenced by synchrony or compensatory dynamics at the population level. We show that variability at both the population and community level is influenced strongly by a few distinct temporal scales: in phytoplankton, ciliate, rotifer, and crustacean communities, synchronous dynamics are predominant at most temporal scales. However, in phytoplankton and crustacean communities, compensatory dynamics occur at a sub-annual scale (and at the annual scale in crustaceans) leading to substantial reductions in community-level variability. Aggregate measures of population and community variability do not detect compensatory dynamics in these communities; thus, resolving their scale dependence unmasks dynamics that are important for community stability in this system. The methods and results presented herein will ultimately lead to a better understanding of how stability is achieved in communities.     

Nitrogen and plankton dynamics in the lagoon of Venice

The nitrogen cycle in the lagoon of Venice, which is the largest Italian lagoon, was investigated by means of a 3D fully coupled transport – water quality model, which had been validated against a substantial amount of real-world data. Nitrogen fluxes among different ecosystem compartments were computed for each month of a reference year, and for each one of the three sub-basins into which the lagoon is conventionally subdivided. The computation included the loads of nitrogen discharged by the tributaries, the direct inputs from the industrial area and the city of Venice, the atmospheric loads, the fluxes at the three lagoon inlets and the internal fluxes between sediment and water compartments and among the three sub-basins. The results of the analysis show that the lagoon, as a whole, exports nitrogen towards the sea. Approximately 4000 tN/year are recycled by the system, while 4640 tN/year is the net input from the drainage basin and the other sources, thus leading to about 8640 tN/year of dissolved inorganic nitrogen that enter the water compartment. Around half of the this amount is used by primary producers, one fourth is exported towards the sea, and one fourth is transferred into the sediment compartment, or lost to atmosphere. These findings suggest that the exchanges through the inlets play an important role in keeping nitrogen concentration at an acceptable level. A more detailed analysis of the model results shows that the non-homogeneous spatial distribution of tributary discharges and point sources is the main cause of the differences in the ecosystem response and water quality among the three sub-basins. Nutrient poorer sub-basins fix a ration of available inorganic nutrient higher than nutrient rich ones. However, they are more efficient in transferring the biomass to the highest trophic levels. Results also include estimates of fluxes that were not quantified so far (such as grazing and recycling), and a validated model, which could have a practical use, for example for assessing implications of reduction of nutrient loads.     

The structure of plankton communities of the Indian Ocean

Species diversity and trophic structure of plankton communities have been considered as most important biological features. The Fisher () and Simpson () indices and the information index H=- pi log₂ pi (Margalef) were used for evaluation of species diversity. The indices have been calculated within the group of filter feeders by numbers ( and H) and biomass ( and H) of the species; the best results were obtained in the latter case. The index H=- pi log₂ pi is proposed for the quantitative evaluation of trophic structure, where a portion of an individual trophic group is taken as pi. The plankton material was collected by the R.V. Vityaz in the northern Indian Ocean in January through February, 1960. Analysis of characteristics of zooplankton distribution, species and trophic diversity in connection with characteristics of the water regime resulted in the definition of 3 types of the plankton communities: (a) those in intensive divergence zones; (b) those in poor divergence zones; (c) those in regions with stable water stratification or with feebly marked convergence, distiguished by different values of biomass, species diversity, and trophic group ratios.     

Study of the nutrient and plankton dynamics in Lake Tanganyika using a reduced-gravity model

An eco-hydrodynamic (ECOH) model is proposed for Lake Tanganyika to study the plankton productivity. The hydrodynamic sub-model solves the non-linear, reduced-gravity equations in which wind is the dominant forcing. The ecological sub-model for the epilimnion comprises nutrients, primary production, phytoplankton biomass and zooplankton biomass. In the absence of significant terrestrial input of nutrients, the nutrient loss is compensated for by seasonal, wind-driven, turbulent entrainment of nutrient-rich hypolimnion water into the epilimnion, which gives rise to high plankton productivity twice in the year, during the transition between two seasons. Model simulations predict well the seasonal contrasts of the measured physical and ecological parameters. Numerical tests indicate that the half saturation constant for grazing by zooplankton and the fish predation rate on zooplankton affect the zooplankton biomass measurably more than that of phytoplankton biomass. This work has implications for the application of this model to predict the climatological biological productivity of Lake Tanganyika.     

Some development patterns of plankton communities in the upwelling areas of the Pacific Ocean

The principal trophic levels, each subdivided into groups of organismic elements, are distinguished in the planktonic communities of the Eastern Equatorial and the Peruvian upwellings. Production intensity or metabolism have been determined experimentally for all elements. A scheme is suggested for computing production from data on metabolism for all the elements of a community, as well as for computing net and real production and other functional characteristics for definite trophic levels and the community as a whole. Based on the quantitative estimation of the efficiency of primary production and other functional characteristics, the development of communities is divided into production and destruction periods; they are, in turn, subdivided into steps associated with a certain degree of water trophicity. The balance of net production of the communities in the Peruvian upwelling indicates that the excess production of a community above the shelf is utilized completely in the narrow (100 to 150 sea miles) band of off-shore water. This paper describes an attempt to trace the changes taking place in the functional characteristics of plankton communities and to compare them with the changes observed in the communities of the Peruvian and East-Equatorial upwellings.     

Simple predator–prey interactions control dynamics in a plankton food web model

A plankton food web model is analysed using interaction parameter values appropriate to the upper mixed layer of the high latitude oceans. The dynamics of this four-variable system are analysed in terms of the dynamics of much simpler two-variable predator–prey subsystems. Thus, the food web's robust, periodic, four-dimensional dynamics are explained by means of two-dimensional spirals and limit cycles. These dynamical subsystems are coupled by means of an omnivore that transfers control of the dynamics between the two predator–prey subsystems. The food web may substantially decouple the predator–prey subsystems so that the oscillating phytoplankton/zooplankton blooms exhibit population collapses when bacterial ‘breathers’ briefly dominate after growing dramatically from low background levels. This regular bloom/breather behaviour becomes benignly chaotic when the system is mildly forced by the annual cycle of the sun's irradiance.     

Cycling of organic nitrogen in marine plankton communities studied in enclosed water columns

Concentrations of fluorescamine-positive substances (primary amines) and turnover rates of L-leucine pools were measured concurrently in seawater samples taken from 1300 m³ plastic enclosures moored in Saanich Inlet, British Columbia, Canada. Concentration and turnover rates of dissolved free amino acids were calculated and then used to determine the instantaneous flux of dissolved free amino acids, which ranged from 0.09 to 2.42 M d^-1 (i.e.,5 to 145 gC l^-1 d^-1). This flux was highest in the euphotic zone, and was related to net primary production but not to the type of dominant primary producer. Comparison of the flux to changes in the concentration of ammonia in deep water suggested that amino acid degradation accounted for 60% of the flux into the ammonia pool. For a given sample, the amino acid carbon flux ranged from 17 to 210% (mean=78%) of the primary production. Such fluxes of amino acid carbon, if used exclusively by the bacterioplankton, would give growth rates ranging from 0.3 to 3.0 (mean=1.7) bacterial doublings d^-1. These calculations indicate that a large fraction of the community carbon and nitrogen flux passes through the bacterioplankton.     

Biomass and production in polar planktonic and sea ice microbial communities: a comparative study

Algal and bacterial biomass and production were measured in the plankton, platelet ice and congelation ice communities at one station in McMurdo Sound, Antarctica during September and October 1986. Bacterial abundances and particulate organic carbon and nitrogen were 10 to 100 times greater in the plankton than in the sea ice, whereas the chlorophyll a concentrations in the plankton and sea ice microbial communities (SIMCO) were similar Rates of both light-limited and light-saturated photosynthesis and daily primary production were 2 to 6 times greater in the plankton than in the SIMCO. Bacterial growth rates ranges from 0.7 to 1.5 d^-1 in all three communities; however, because of the greater bacterial biomass in the plankton, bacterial production was 15 to 20 times higher there than in the SIMCO. These results suggest that during the early austral spring, planktonic production contributes significantly to total production in ice-covered environments.     

A simulation model of carbon and oxygen dynamics in a reservoir

The CORES model represents the long-term carbon and oxygen dynamics of the Conowingo Reservoir. The model, formulated using subsetted historical data, is readily validated using the remaining historical data for all state variables of importance. Sensitivity analyses make it clear that the flux of particulate organic carbon from upstream sources and its subsequent “trapping” and metabolism in the deeper regions of the reservoir are the major factors controlling the oxygen declines historically observed in the summer months. Increased natural vertical mixing (i.e., transport of oxygen from shallow to deep waters) can increase deep-water oxygen concentrations to some extent, but not to the degree necessary to meet water quality standards. Air-to-water diffusion adds substantial quantities of oxygen to surface waters in these mixing events. Simple solutions, such as requiring continuous flow, do not appear to adequately increase the dissolved oxygen levels in the deeper portion of the reservoir.     

Development of north sea coastal plankton communities in separate plastic bags under identical conditions

In two experiments lasting 4 to 6 weeks, communities of North Sea coastal plankton kept in separate plastic bags (of about 1400 l) and exposed to the same environmental conditions showed very similar patterns of growth and decline. This result means that the method is suitable for the evaluation of toxic effects of environmental pollutants at low concentrations on complex plankton systems. The phytoplankton in the bags produced a succession of blooms, which were probably limited by shortage of nutrients. The dominant zooplankton organisms were various species of copepods which can develop in the bags from egg to adult. Strong indications were found that mineralization of organic matter occurs in the bags. Chemical parameters and phytoplankton biomass were found not to be stratified, indicating that the contents of the bags were well mixed.Work carried out under Contract No. 110-75-1 ENVN of the E.C. Environmental Research Programme.     

Production and respiration of overall plankton and ultraplankton communities at the entrance to the Gulf of Finland in the Baltic Sea

Primary productivity and respiration of the overall plankton community and of ultraplankton (organisms passing through a 3-m Nuclepore filter) were studied at the entrance to the Gulf of Finland during the growth season in 1982. Data of the respiration measurements from previous years are also presented. During the development of a diatom spring bloom, the algal component could be successfully separated from the bacterial component by size fractionation with a 3-m Nuclepore filter and thus the algal respiration could be approximated, being on the order of 10 to 20% of the gross production. After the phytoplankton spring maximum, bacteria played an important role in mediating the energy flow from phytoplankton exudates to higher trophic levels. Maximum values of 1 230 and 740 mg O₂ m^-2 d^-1 were recorded for overall and for ultraplankton respiration, respectively, during late July. High productivity values coupled with low phytoplankton biomass and low inorganic nutrient values were also recorded in late July, indicating effective nutrient regeneration and rapid turnover of the plankton community. During late summer, a considerable fraction (over 30%) of phytoplankton production was released as exudates, suggesting that much of the energy is channeled to higher trophic levels via bacterial pathways rather than by direct herbivorous grazing during this season. The summer development of phytoplankton community structure and functioning is strongly controlled by hydrographic conditions, i.e. by nutrient inputs via upwelling and by water temperature. A carbon budget for late summer indicated that bacteria may contribute only up to 50% of the overall respiration of the plankton community, which suggests that heterotrophs other than bacteria play an important role in nutrient regeneration. The present study stresses the importance of energy flow via the phytoplankton exudatebacteria-micrograzer pathway in relatively oligotrophic, brackish water ecosystems.     

Origins and scales of hypoxia on the Louisiana shelf: Importance of seasonal plankton dynamics and river nutrients and discharge

Management plans for the Mississippi River Basin call for reductions in nutrient concentrations up to 40% or more to reduce hypoxia in the Gulf of Mexico (GOM), while at the same time the government is considering new farm subsidies to promote development of biofuels from corn. Thus there are possibilities of both increasing and decreasing river nutrients depending on national priorities. River flow rates which also influence the extent of hypoxia on the shelf may be altered by global climate change. We have therefore developed a series of simulations to forecast ecosystem response to alterations in nutrient loading and river flow. We simulate ecosystem response and hypoxia events using a linked model consisting of multiple phytoplankton groups competing for nitrogen, phosphorus and light, zooplankton grazing that is influenced by prey edibility and stoichiometry, sub-pycnocline water-column metabolism that is influenced by sinking fecal pellets and algal cells, and multi-element sediment diagenesis. This model formulation depicts four areas of increasing salinity moving westward away from the Mississippi River point of discharge, where the surface mixed layer, four bottom layers and underlying sediments are represented in each area. The model supports the contention that a 40% decrease in river nutrient will substantially reduce the duration and areal extent of hypoxia on the shelf. But it also suggests that in low and middle salinity areas the hypoxia response is saturated with respect to nutrients, and that in high salinity regions small increases in nutrient and river flow will have disproportionally large effects on GOM hypoxia. The model simulations also suggest that river discharge is a stronger factor influencing hypoxia than river nutrients in the Mississippi River plume. Finally, the model simulations suggest that primary production in the low salinity regions is light limited while primary production in the higher salinity zones is phosphate limited during the May to October period when hypoxia is prevalent in the Mississippi River plume.     

Autotrophic and bacterial production: Fractionation of plankton populations by differential filtration of samples from the English channel

The relationship between release of organic substances by phytoplankton and their utilization by heterotrophic bacteria has been measured by means of differential filtration in parallel Steemann Nielsen and Parsons and Strickland incubations. In Southampton Waters (UK), between March and September, the values for bacterial production measured in the above manner varied between 1 and 30%, the total primary production being taken as 100%. Even when low bacterial production occurs during the light period of incubation, a significant increase takes place during the following dark period. Thus, a close correlation between primary algal and the resultant bacterial production can be demonstrated. Quantitative data on bacterial production may be obtained from the conventional productivity experiments by simple double filtration.     

Effect of nitrogen source on short-term light and dark CO2 uptake by a marine diatom

Based on a series of short-term incubations involving the marine diatom Chaetoceros simplex (Bbsm), precultured in NH ₄ ⁺ -, NO ₃ ^- -and urea-limited continuous cultures at several dilution rates, we found that both the short-term specific rate of ¹⁴CO₂ uptake and the amount of CO₂ fixed after 8- and 16-min incubations were unaffected by enrichment with NH ₄ ⁺ , urea, or NO ₃ ^- when NH ₄ ⁺ or urea were the preconditioning forms of N, but were slightly suppressed when the cells were first grown on NO ₃ ^- . Similar enrichments in the dark, however, led to significant CO₂ uptake under all conditions of NH ₄ ⁺ enrichment and to similarly enhanced CO₂ uptake, but only at high growth rates, when urea was the source of enrichment nitrogen. Our light results are contrary to some contemporary findings, but there does seem to be agreement that photosynthetic rates of rapidly growing phytoplankton will not be affected by exposure to pulses of nitrogen. Enhanced dark uptake, in contrast, appears to be characteristic of phytoplankton under all degrees of N limitation, and, as such, may be useful as an “all or nothing” index of the nitrogen status of natural waters. There is some indication that the index may be useful in determining both the form of and the degree of N limitation as well.     

Microprofiles of oxygen in epiphyte communities on submerged macrophytes

Mussels (Mytilus edulis) transferred in net bags from clean to chronically mercury polluted water readily accumulated mercury during an exposure period of three months. Growth of the transplanted mussels had a diluting effect on the mercury concentration, but the absolute weight of mercury uptake increased throughout the entire period, though there was a tendency for decreased efficiency of the removal of mercury per liter of water filtered by the mussels. Mussels were also translocated from polluted to clean (laboratory) water to depurate mercury. The biological half-lives of mercury was 293 d for M. edulis from the chronically polluted area in contrast to only 53 d for mussels from a temporary massive mercury polluted area near a chemical deposit. In both cases about 75% of the total mercury in the mussels was inorganic, and it is suggested that both inorganic and organic mercury species were immobilized in mussels from the long-term mercury polluted area, whereas the immobilization capacity was exceeded in the short-term mercury exposed mussels near the chemical deposit. Very slow elimination of mercury was observed in the deposit-feeding bivalve Macoma balthica from the chronically polluted area, and about 6% of the total mercury was methyl-+phenyl-mercury. This is more than three times lower than found in M. edulis from the same collecting site. A pronounced difference in the mercury speciation (i.e., total mercury, total organic mercury, methyl-mercury and phenyl-mercury) in M. edulis from the two mercury polluted areas is thought to reflect the different character of the mercury pollution in the two areas.     

Temporal dynamics of arsenic uptake and distribution: food and water risks in the Bengal basin

Abstract

Contaminated food chain is a serious contender for arsenic (As) uptake around the globe. In Nadia, West Bengal, we trace possible means of transfer of As from multiple sources reaching different trophic levels, and associated seasonal variability leading to chronic As uptake. This work considers possible sources-pathways of As transfer through food chain in rural community. Arsenic concentration in groundwater, soil, rice, and vegetable-samples collected detected in different harvest seasons of 2014 and 2016. Arsenic level in shallow groundwater samples ranged from 0.1 to 354?µg/L, with 75% of the sites above the prescribed limit by WHO (10?µg/L) during the boro harvest season. High soil As content (～20.6?mg/kg), resulted in accumulation of As in food crops. A positive correlation in As conc. with increase over period in all sites indicating gradual As accumulation in topsoil. Unpolished rice samples showed high As content (～1.75?mg/kg), polishing reduced 80% of As. Among vegetables, the plant family Poaceae with high irrigation requirements and Solanaceae retaining high moisture, have the highest levels of As. Contaminated animal fodder (Poaceae) and turf water for cattle are shown to contaminate milk (0.06 to 0.24?µg/L) and behoves strategies, practices to minimize As exposure.     

Predation,recruitment and the dynamics of communities of coral-reef fishes

I investigated the ability of predators to influence the patterns of species richness and abundance of non-piscivorous fishes on small, artificial reefs replenished by natural recruitment. Periodic removal of predators effectively reduced the species richness and abundance of predators on removal reefs. The difference between the number of predators on control and removal reefs was greatest immediately following the removal of predators and attenuated between removals. During periods of recruitment, species richness and total abundance of recently-recruited, non-piscivorous fishes were generally greater on predator-removal reefs than on control reefs. Species richness and total abundance of resident non-piscivorous fishes were not affected by the removal of predators in the first year of the experiment. Both abundance and species richness of residents, however, were greater on the removal reefs during the second year of the experiment. The difference in the responses of the two age classes to the removal of predators suggests that predators may affect community patterns of older age classes through time-lagged effects on the survivorship of younger age classes. At the end of the experiment, species richness was positively related to abundance for recruits and residents. The effects of removing piscivorous fishes on the abundance of non-piscivorous fishes were similar for species considered separately. A greater number of species of recruit and resident fishes were more abundant on reefs from which predators had been removed. These data suggest that predators can play an important role in structuring communities of fishes on coral reefs.     

Macroinfauna of a Southern New England salt marsh: seasonal dynamics and production

The animal-habitat relationships and seasonal dynamics of the benthic macroinfauna were investigated from November 1986 to October 1988 in the Great Sippe-wissett salt marsh (Massachusetts, USA). Total macrofaunal abundance varied seasonally, displaying a peak in late spring and early summer, then declining sharply during late summer and recovering briefly in fall before collapsing in winter. Three macroinfaunal assemblages were found in the marsh, distributed along gradients of environmental factors. These included a sandy non-organic sediment assemblage, a sandy organic sediment assemblage and a muddy sediment assemblage. The species groups characteristic of unstable sandy non-organic sediments included the polychaetes Leitoscoloplos fragilis, Aricidea jefreyssi, Magelona rosea and Streptosyllis verrilli, the oligochaete Paranais litoralis, and the crustacean Acanthohaustorius millsi. Sandy organic sediments were characterized by the polychaetes Marenzelleria viridis, Capitella capitata, Neanthes succinea, N. arenaceodonta, Polydora ligni and Heteromastus filiformis, the oligochaete Lumbricillus sp., and the mollusc Gemma gemma. In muddy sites, the polychaete Streblospio benedicti and the oligochaetes Paranais litoralis and Monopylephorus evertus were the dominant species. Secondary production of benthic macroinfauna in each of these habitats was estimated. The highest values of biomass and production were recorded in the sandy organic sediments. Secondary production was estimated to be 1850 kJ m^-2 yr^-1 in sandy organic areas, but only 281 kJ m^-2 yr^-1 in sandy non-organic areas and 113 kJ m^-2 yr^-1 in muddy areas. This results in an area-weighted average production of 505 kJ m^-2 yr^-1 for the unvegetated areas of the marsh. The Great Sippewissett salt marsh has an area of 483800 m², the total secondary production of the macroinfauna for the whole unvegetated area of the marsh was estimated as 4651 kg dry wt yr^-1, expressed as somatic growth. This production value seems consistent with production data obtained for other intertidal North Atlantic environments.     

Composition,structure and productivity of neritic plankton communities near the bird colonies of the northern shores of Novaya Zemlya

A survey was made of coastal waters around the colonies of Arctic sea birds, which showed that such zones are highly productive. The mean biomass (standing stock) of crustacean plankton for the whole water column varied from 470 to 670 mg/m³ in separate transects, its maxium value reaching 1950 mg/m³. A list of species is given. Dominance hierarchy among the species is clearly expressed. The stable discrete structure of the community is assured by the spatial disjunction of dominant-form populations and the existence of the latter as relatively isolated aggregations. The conclusions arrived at are based on the materials of net hauls and analyses of the food of planktophageous birds (Plotus alle, family Alcidae). Favourable hydrological factors in conjunction with the fertilization of water with bird guano assure the stability of this neritic community with its multiplicity of interrelationships between producers and consumers of the first and second trophic levels, existing under the rigorous conditions of an ice-covered region.     

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.