The effects of seston food quality on planktonic food web patterns

In planktonic food webs, the conversion rate of plant material to herbivore biomass is determined by a variety of factors such as seston biochemical/elemental composition, phytoplankton cell morphology, and colony architecture. Despite the overwhelming heterogeneity characterizing the plant–animal interface, plankton population models usually misrepresent the food quality constraints imposed on zooplankton growth. In this study, we reformulate the zooplankton grazing term to include seston food quality effects on zooplankton assimilation efficiency and examine its ramifications on system stability. Using different phytoplankton parameterizations with regards to growth strategies, light requirements, sinking rates, and food quality, we examined the dynamics induced in planktonic systems under varying zooplankton mortality/fish predation, light conditions, nutrient availability, and detritus food quality levels. In general, our analysis suggests that high food quality tends to stabilize the planktonic systems, whereas unforced oscillations (limit cycles) emerge with lower seston food quality. For a given phytoplankton specification and resource availability, the amplitude of the plankton oscillations is primarily modulated from zooplankton mortality and secondarily from the nutritional quality of the alternative food source (i.e., detritus). When the phytoplankton community is parameterized as a cyanobacterium-like species, conditions of high nutrient availability combined with high zooplankton mortality led to phytoplankton biomass accumulation, whereas a diatom-like parameterization resulted in relatively low phytoplankton to zooplankton biomass ratios highlighting the notion that high phytoplankton food quality allows the zooplankton community to sustain relatively high biomass and to suppress phytoplankton biomass to low levels. During nutrient and light enrichment conditions, both phytoplankton and detritus food quality determine the extent of the limit cycle region, whereas high algal food quality increases system resilience by shifting the oscillatory region towards lower light attenuation levels. Detritus food quality seems to regulate the amplitude of the dynamic oscillations following enrichment, when algal food quality is low. These results highlight the profitability of the alternative food sources for the grazer as an important predictor for the dynamic behavior of primary producer–grazer interactions in nature.     

Polonium-210 and lead-210 in marine food chains

²¹⁰Po and ²¹⁰Pb have been measured systematically in whole animals, muscle and hepatopancreas of crustaceans and of molluscan cephalopods representative of a pelagic and benthic food chain. The same nuclides were also measured in liver, pyloric caecum, stomach contents and muscle of tuna. The concentration factors from sea water to whole animals were approximately constant along both food chains, being of the order of 10⁴ for ²¹⁰Po and 10² for ²¹⁰Pb. The highest concentration factors were found in shrimp of the genus Sergestes. In muscle, the concentration factors were an order of magnitude less; in the hepatopancreas, they were an order of magnitude higher, reaching 10⁶ in shrimp of the genus Sergestes. Such concentrations imply alpha-radiation doses of the order of 10 rem per year and more in this organ, which contains about 50 to 90% of the ²¹⁰Po in the whole animal in the 11 species analyzed. A detailed study of the intracellular behaviour of ²¹⁰Po in the hepatopancreas is clearly indicated. ²¹⁰Po can be used as a sensitive natural tracer in biological systems. Thus, feeding Meganyctiphanes norvegica in the laboratory on food low in ²¹⁰Po led to an approximate value of about 61/2 days for the biological half-life of ²¹⁰Po in the hepatopancreas of this euphausiid. Furthermore, the data on ²¹⁰Po and ²¹⁰Pb in the cephalopod hepatopancreas allowed the time of conservation of frozen squid which had been bought at the market to be estimated.     

A directly coupled,artificial two-step food chain for long-term experiments with filter-feeders at constant food concentrations

A system is described consisting of a chemostat for culturing algae, a dilution unit and special experimental vessels, allowing long-term experiments with filter feeders at a constant concentration and quality of food. The algal suspension flows continuously from a chemostat, is mixed with sterile filtered water, and pumped directly into the experimental vessels. The flow through the vessels is intensive enough to avoid a significant change in food concentration by sedimentation or grazing. There are two possible arrangements, the first providing many duplicates with exactly the same food concentrations, the second providing a series of graduated food concentrations. Details of construction are given. The application of both arrangements is demonstrated by experiments with Daphnia pulex. Further possibilities of application are suggested.     

Food ingestion by the marine planktonic copepod Paracalanus in relation to abundance and size distribution of food

Juveniles and adult females were presented a food spectrum of three algae of different sizes (4.5, 12 and 20 m cell width). The increase in rate of ingestion of the medium-sized alga with an increase in copepod size was significantly greater than the increase in rate of ingestion of the small alga. It is hypothesized that the perception of chemical signals from the small alga by a copepod decreases as the copepod moults from stage to stage. The rate of ingestion of the large alga by copepod stage V (CV) and adult females was lower than the rate of ingestion of the medium-sized alga at mid- and high phytoplankton concentrations. The amount of nitrogen ingested when the medium-sized alga alone was offered was either higher than (stage C II) or not significantly different from that when the three algae were offered together (stage C IV). Ingestion rates are reduced when there is a multialgal food source. This implies that there is increased stability in the ocean because multiparticle food sources are more slowly depleted than unialgal foods. Weight-specific ingestion rates of copepods fed the three algae simultaneously increased from nauplius to stage C III and then decreased as adulthood approached. The contribution of the small alga to the total amount of nitrogen ingesied was greatest for naupliar stages while the contribution of the medium-sized cells was greater for later stages. The largest alga was readily ingested by stage C V and adult females but never contributed more than 25% of the nitrogen ingested. Eight to 12% of the nitrogen ingested by adult females was from the small alga. It is hypothesized that the algal cell size for maximum nitrogen ingestion in upwelled waters is relatively small, round or square and close to the size threshold below which adult females do not sense individual cells.     

Variable transfer of detoxified metals from snails to hermit crabs in marine food chains

In any polluted marine environment, different invertebrate species contain markedly different concentrations of heavy metals. Primary producers take metals from seawater, but animals take additional metals from diets of animals, plants and detritus. Metals in a dietary organism of a food chain have varying reactivities and they follow different biochemical pathways. Excess metals are bound by ligands to form insoluble compounds within cytological compartments. These metabolic systems prevent the disruption of normal biochemical reactions by metals. The present work on Mediterranean invertebrates, initiated in Greece in 1993, used digestive glands from three species of marine snail,Monodonta mutabilis (Philippi),Cerithium vulgatum (Bruguière), andMurex trunculus (Linnaeus) as prey tissue, and hermit crabsClibanarius erythropus (Latreille) as predators; the digestive glands and faecal pellets from all animals were analysed by atomic absorption spectroscopy and x-ray microanalysis. Most metals detoxified by the snails are unavailable to the crabs and they pass straight through the gut and appear in the faecal pellets. This applies to significant proportions of the manganese, nickel, copper, zinc and silver which are bound electrostatically to phosphate or covalently to sulphur within membrane-bound intracellular compartments. Cadmium and chromium are transferred to the crabs. In digestive glands of snails, cadmium is bound to soluble highsulphur protein in the cytosol; the cytology of chromium in these animals is not known.     

Flock size,food dispersion,and the feeding behavior of crossbills

Summary Feeding rates of five captive red crossbills (Loxia curvirostra) were measured when they were foraging alone, and in flocks of two or four on three seed dispersion patterns. On the most strongly clumped seed dispersion, individuals had higher mean feeding rates and the smallest probability of starvation when in flocks of two than when alone or in flocks of four. Individuals in flocks of four had higher feeding rates on the weakly clumped seed dispersion than on the uniform and more clumped seed dispersions; there were no food finding benefits gained from flocking on the uniform seed dispersion and aggression increased as food became more clumped. Most recent work has assumed that flocking results in higher feeding rates only because time spent vigilant is reduced. Crossbills, however, did not visit more cones per unit time as flock size increased, as would be expected if less time was spent vigilant. Thus, any reductions in vigilance as flock size increased were countered by increases in other behaviours, such as those related to aggression. Consequently, the higher mean feeding rates of crossbills in flocks than when solitary is not attributable to reduced vigilance. The increase in mean and the decline in variance of feeding rates occurred because crossbills in flocks found good patches earlier, and possible by spending less time assessing poor patches.     

The use of radioactive isotopes to measure the transfer of materials in aquatic food chains

Radioisotopes have been misused extensively by ecologists in transfer studies within food chains. Unless it is known that no recycling of isotope has occurred during the experiment, the assumption of linear uptake when in fact the system is not linear, even over short periods, can lead to significant errors in the estimation of ingestion or feeding. If recycling occurs, at least a 3 or 4-compartment, hydraulic-type model is necessary to even approximate the complicated kinetics of isotopic transfer in a simple aquatic feeding experiment. In any event, it is essential to follow the uptake or loss of an isotope (change in specific activity) as a function of time in at least 1 compartment before deciding on an appropriate model. If experiments are designed so that the maximum number of rate processes are summed or integrated by the animal, the kinetics can be considerably simplified. If the food supply is uniformly labelled, the rate of change of tracer can be used to give a rate of ingestion (grazing). If the predator is labelled with a suitable isotope before starting the experiment, the rate of loss of its isotope burden under different experimental conditions can be used to determine respiration or excretion rates, turnover rates, ingestion, and the size and number of major compartments in the transfer system.Bedford Institute Contribution. Woods Hole Oceanographic Institution Contribution No. 2690.     

Anthropogenic subsidies alter stream consumer-resource stoichiometry, biodiversity, and food chains.

Urbanization is dramatically changing nutrient and organic matter regimes in streams, yet the community and ecosystem implications often remain obscure. We assessed the consequences of sewage-derived particulate organic matter (SDPOM) for invertebrate community structure and function in a headwater stream. Using stable isotope analyses, we found assimilation of organic SDPOM to double community secondary production, and stoichiometric analyses revealed SDPOM enriched in phosphorus (P) to foster putatively fast-growing, P-rich consumers in the subsidized reach. This altered consumer-resource stoichiometry impacted both community structure and nutrient fluxes through the invertebrate community. Community structure shifted toward significantly reduced diversity and evenness in the subsidized reach and consequently toward shorter food chains. Our integration of ecological stoichiometry with stable isotope analyses and food web ecology expands the previous focus of traditional ecotoxicology and ecophysiology to an ecosystem-level appreciation of pollutant ecology.     

Arsenic trophodynamics along the food chains/webs of different ecosystems: a review

In drinking water, arsenic (As) food chain accumulation may pose a risk to human health. An attempt was made to synthesise the published information concerning As trophic transfer along the food chain/web of various marine, freshwater, and terrestrial ecosystems. Some investigations included As speciation. Further objectives were to outline the factors potentially influencing As trophodynamics and to understand the consequence of As trophic transfer in the environment.     

Nitrogen cycling in a marine planktonic food chain: Nitrogen fluxes through the principal components and the effects of adding copper

Nitrogen cycling in a natural planktonic food chain was followed before and after perturbation by copper (10 g l^-1). Observations were separated into effects from artificial containment of the food chain and effects from the addition of copper. An initial reduction of seston-N and subsequent recovery, uniformly high settling rates of suspended particulate-N, and general decrease in zooplankton-N were attributed to the physical limitations imposed by the enclosures (e.g. elimination of normal mixing processes). Acute inhibition of nitrate assimilation by the seston, acute elevation of seston-N settling and chronic inhibition of zooplankton feeding, along with accelerated decline in zooplankton stocks, were attributed to copper stress. Since zooplankton stocks and fluxes were most drastically affected by copper, its overall effect on N-cycling through the planktonic food chain would depend largely upon the importance of zooplankton in regulating the production of the remaining seston.     

Comparison of mercury bioaccumulation between wild and mariculture food chains from a subtropical bay of Southern China

Bioaccumulation and trophic transfer of mercury (Hg) both in the natural marine ecosystem and the mariculture ecosystem were studied at Daya Bay, a subtropical bay in Southern China. Averaged Hg concentrations in sediment, phytoplankton, macrophyte, shrimp, crab, shellfish, planktivorous fish, carnivorous fish, farmed pompano, farmed snapper, compound feed and trash fish were 0.074, 0.054, 0.044, 0.098, 0.116, 0.171, 0.088, 0.121, 0.210, 0.125, 0.038 and 0.106 μg g⁻¹ dw, respectively. These Hg levels were at the low–middle ends of the global range. Positive correlation between Hg concentrations in farmed fish and fish weights/sizes was observed, whereas no clear correlation between Hg concentrations and lipid contents was found. Hg concentrations followed macrophyte < phytoplankton < sediment < planktivorous fish < shrimp < crab < carnivorous fish < shellfish, and commercial feed < trash fish < farmed fish. Hg was biomagnified along the marine food chain in the ecosystem of Daya Bay. Hg levels in the farmed fish were higher than those in the wild fish primarily because of the higher Hg level in fish feed and the smaller size of marine wild fish.

     

Particulate DNA in subtropical oceanic and estuarine planktonic environments

Particulate DNA was measured in estuarine, coastal, and oligotrophic oceanic environments near the southwest coast of Florida and in the Gulf of Mexico. Particulate organic carbon and nitrogen (POC and PON), chlorophyll a, bacterial direct counts, DNA, and bacterial activity as determined by thymidine incorporation all showed a high degree of intercorrelation. Normalization of the data for offshore/onshore similarities by dividing by POC yielded significant correlations only for DNA, direct counts, and bacterial activity. Most (70–99%) of the particulate DNA in offshore samples was in the 0.2- to 1-m fraction, while DNA in nearshore and estuarine samples was associated with larger particles. Cellular DNA contents obtained by dividing DNA by direct counts in the 0.2- to 1-m fraction were in the range of reported bacterial genome weights. However, DNA nitrogen comprised a greater proportion of the PON than reported for microorganisms in culture. Collectively, these results suggest that (1) most of the particulate DNA in oceanic environments is contained in bacterioplankton; (2) DNA is a significant proportion of the cell biomass, possibly due to growth under nutrientlimiting conditions.     

Respiration,photosynthesis and carbon metabolism in planktonic ciliates

Release of¹⁴C-labelled carbon dioxide from uniformly labelled cells was used to measure respiration by individual ciliates in 2-h incubations in 1989 and 1990. In a strictly heterotrophic ciliate,Strobilidium spiralis (Leegaard, 1915), release of labelled carbon dioxide was equivalent to ca. 2.8% of cell C h^–1 at 20°C, and there was no difference between rates in the dark and light. In the chloroplast-retaining ciliatesLaboea strobila Lohmann, 1908,Strombidium conicum (Lohmann, 1908) Wulff, 1919 andStrombidium capitatum (Leegaard, 1915) Kahl, 1932, release of labelled carbon dioxide was less in the light than in the dark in experiments done at 15°C. InL. strobila release of radiolabel as carbon dioxide was equivalent to ca. 2.4% of cell C h^–1 in the dark but ca. 1% at 50µE m^–2 s^–1, an irradiance limiting to photosynthesis. InS. conicum release of radiolabel as carbon dioxide was equivalent to ca. 4.4% of cell C h^–1 in the dark, but at an irradiance saturating to photosynthesis (250 to 300µE m^–2 s^–1) there was no detectable release of labelled carbon dioxide. InS. capitatum release of radiolabel as carbon dioxide was equivalent to ca. 4.3% of cell C h^–1 in the dark but at an irradiance saturating to photosynthesis was ca. 2.4% of cell C h^–1. These data, combined with data from photosynthetic uptake experiments, indicate that¹⁴C uptake underestimates the total benefit of photosynthesis by 50% or more in chloroplastretaining ciliates.Contribution no. 7510 from the Woods Hole Oceanographic Institution     

Kinetics of light-intensity adaptation in a marine planktonic diatom

The marine planktonic diatom Thalassiosira weisflogii was grown in turbidostat culture under both continuous and 12 hL: 12 hD illumination regimes in order to study the kinetics of adaptation to growth-irradiance levels. In both illumination regimes adaptation to a higher growth-irradiance level was accompanied by an increase in cell division rates and a decrease in chlorophyll a cell^-1. The rates of adaptation for both processes, derived from first order kinetic analysis, equaled each other in each experiment. The results suggest that during the transition from low-to-high growth-irradiance levels chlorophyll a is diluted by cell division and is not actively degraded. Introduction of a light/dark cycle lowered the rate of adaptation. In transitions from high-to-low growth-irradiance levels there was a sharp drop in growth rates and a slow increase in chlorophyll a cell^-1 under both continuous and intermittent illumination. In the 12 hL:12hD cycle there was a circadian rhythm in chlorophyll a cell^-1, where cellular chlorophyll contents increased during the light cycle and decreased during the dark cycle. This circadian rhythm was distinctly different from light intensity adaptation. For kinetic analysis of light intensity adaptation in a 12 hL: 12 hD cycle, the circadian periodicity was separated from the light intensity response by subjecting the data to a Kaiser window optimization digital filter. Kinetic parameters for light-intensity adaptation were resolved from the filtered data. The kinetics of lightintensity adaptation of marine phytoplankton are discussed in relation to their spatial variations and time scales of mixing.This research was performed at Brookhaven National Laboratory under the auspices of the United States Department of Energy under Contract No. DE-AC02-76 CH00016     

Ontogenic changes of amino acid composition in planktonic crustacean species

Changes in amino acid composition (AAC) during ontogeny of some planktonic crustacean species commonly found in fresh and brackish coastal waters were compared. For these comparisons two calanoid copepods (Eurytemora velox and Calanipeda aquae-dulcis), two cyclopoid copepods (Diacyclops bicuspidatus odessanus and Acanthocyclops robustus) and two Daphnia (Daphnia pulicaria and Daphnia magna) species were selected. A discriminant analysis was performed to determine whether there were significant differences between the AAC of the different stages of each species. Results show gradual changes in AAC during ontogeny of the copepod species. Calanoids showed the greatest differences in AAC between stages, followed by cyclopoids. Gradual changes in AAC were due to the increase in some amino acids such as alanine, valine, glutamic acid, glycine, arginine, proline and tyrosine from nauplii to adults. The latter showed a remarkable increase in all copepod species. In contrast, Daphnia species showed a relatively constant AAC during development, with only minor changes being detected, and not related with ontogeny. Differences in the physico-chemical variables of the lagoons do not seem to be the cause of copepod ontogenic changes in AAC. Data suggest that AAC differences found between stages of copepod species could indicate a gradual change in diet during the life cycle of these copepods.     

Diel patterns of planktonic bioluminescence in the northern Sargasso Sea

Day-night changes in the vertical distribution, intensity, and size of bioluminescence flashes were investigated during a series of cruises to the northern Sargasso Sea in 1987 and 1988. Overall, depth integrated bioluminescence potential and flash density estimated from in situ measurements with a pumping bathyphotometer were 2 to 5 x higher at midnight than at midday. Depths from 50 to 100 m exhibited the most substantial day to night increases in bioluminescence potential and flash density. When classified by flash size (photon output per flash event), the increase from day to night was significant for all flash sizes, but was most dramatic for small flashes producing <7 x 10⁸ photons flash^-1. Bioluminescence potential and flash density increased 2 to 3 x during bathyphotometer measurements made at dusk. Bioluminescent light budgets estimated from day and night net collections in May and August 1987 also predicted 2.5 x higher nighttime than daytime mesoplankton bioluminescence. However, large bioluminescent taxa (mesoplankton) capable of significant vertical migrations only contributed on the order of 15% of the total bioluminescence in surface waters. Our results do not support the idea that most of the nightly increase in bioluminescence potential and flash density are due to vertical migration of bioluminescent organisms; rather they are consistent with an alternate view that photoinhibition of bioluminescent flashing by dinoflagellates may be primarily responsible for the diel patterns.     

Modelling the effect of constant and fluctuating food supply on egg production rates of Acartia grani

A mathematical model of the individual budget of a spawning female of the copepod Acartia grani (Sars) has been used to simulate the time-scale of egg production over various external forcings (or inputs) of food fluctuation conditions. The budget matter in the body of the copepod females is distributed through four compartments: the whole digestive tract (globally named as gut), the hemolymph (which include the body fluid with available nutriments for the organs), the structural body weight, and the gonad. This small calanoid species does not carry lipid reserves but cumulate some labile reserves in its body, according to food availability. The model results show how the continuous spawning varies with food fluctuations, and suggest the mechanisms inducing the delay of response to starvation by using the metabolic reserves. Three different patterns in egg production response are observed: food fluctuations with frequencies below 12 h have no effect on egg production; food fluctuations of 12 h to 5 days induce synchronous egg production fluctuations; beyond 5 days the strong physiological changes induced by long starvation durations create delays in the responses to food replenishment. The available data of cultivated cohorts under laboratory conditions are used to validate the model. The properties underlined by the model, in particular its weak capacity to respond to starvation, allow explaining A. grani distribution in specific habitats. Different experimental protocols for complementary experiments are proposed to complete the model validation in other forcing conditions.     

Pools of chlorophyll and live planktonic diatoms in aphotic marine sediments

Chlorophyll a and numbers of live pelagic diatoms were recorded from sediment depth profiles at 11 stations in the oligotrophic Øresund, Denmark, in late-June. Extraction efficiency of chlorophyll a analysed fluorometrically did not differ significantly between paired samples of frozen-thawed and fresh sediment. The depth profiles of chlorophyll a could be explained by a diagenetic model involving two different chlorophyll pools: one reactive pool declining exponentially with core depth, and one non-reactive pool, of about 1 µg Chl ml^-1 wet sediment, being constant with depth. The number of live diatoms, quantified by the dilution-extinction method, and expressed in terms of most probable number (MPN), declined from an average of about 300,000 g^-1 in the surface sediment to zero values at a depth of 13 cm. The number of live cells was significantly correlated with the sediment chlorophyll a, and the profiles of live cells as well as reactive chlorophyll followed the same exponential decline with core depth, suggesting that the main source of chlorophyll in the sediment was live pelagic diatoms. Taxonomic composition of diatoms in the sediment, dominated by the pelagic genera Chaetoceros, Thalassiosira and Skeletonema, matched the species composition in the water column 3 months earlier during the spring bloom. Regular recordings of the phytoplankton community in the water column showed that only these specific bloom species could be the source of the sediment content of diatoms and chlorophyll a. Further, the ratios between live cells and chlorophyll a were similar in the sediment and in the spring bloom. A conservative estimate of depth-integrated pools of diatoms in the sediment suggested that about 44% of the total phytoplankton biomass during the spring bloom was still present as live cells in the sediment after 3 months. This indicates that the spring bloom input to the sediment is not degraded immediately by the benthic fauna.     

Microgeographical shell variation in Littorina striata, a planktonic developing periwinkle

Littorina striata is a strictly Macaronesian, intertidal periwinkle with planktonic development. The species produces both nodulose and smooth shells, which co-occur at Ilheu de Vila Franca do Campo, a drowned crater situated about 1000?m off the south coast of São Miguel, Azores. The present work describes and analyzes the shell variation, temporal change and ecological distribution of the two shell types at this crater over a 3-year period. Nodulose shells were more common in the sheltered lagoon inside the crater, while smooth specimens dominated the outside of the wave-exposed crater. Moreover, nodulose specimens were smaller and weighed less compared to smooth ones. However, regardless of morphotype, shells from the lagoon had a smaller aperture and were less globose than those from the outside. Within an exposure regime, smooth specimens had a larger aperture. These patterns remained constant over time. They are tentatively interpreted as functional adaptations to thermal stress and wave exposure. The mechanisms that maintain these patterns are still unknown. Yet, aperture height of specimens transplanted from the lagoon to the outside increased markedly over a period of 5 months, whereas no comparable changes were observed in other experimental groups (i.e. transplanted from the outside into the lagoon, transplanted within the outside and transplanted within the lagoon; the latter two transplant groups being the blancos). This suggests that at least the aperture size in L. striata may be an ecophenotypically plastic trait.     

Giant larvae: Prolonged planktonic larval phase in the asteroidLuidia sarsi

The bipinnaria larva of the shelf-dwelling asteroidLuidia sarsi Duben and Koren, 1845, can attain (exceptionally) a length of 35 mm; metamorphosis to the adult benthic phase usually occurs well before this size is reached. A seastar rudiment develops at the oral end of the bipinnaria and, under normal circumstances, it detaches itself from its larval body before reaching a diameter of 5 mm. Midwater trawl samples taken from the surface to 1650 m in the Porcupine Seabight during 1978 and 1979, however, includedL. sarsi larvae with post-larval rudiments of 15 mm diameter and greater. It is shown that the bipinnariae and attached post-larvae both continued to grow until a point was reached when it appeared that the young sea-star could no longer be sustained by the bipinnaria. There is evidence that the largest post-larvae had absorbed the bipinnaria tissue and this, it is concluded, is the final stage before enforced settlement. From the estimated growth rate it is likely that many specimens had been in the plankton for >1 yr, giving this species a capability for long-distance dispersal.