Stoichiometry and food-chain dynamics

Traditional models of chemostat systems looking at interactions between predator, prey and nutrients have used only a single currency, such as energy or nitrogen. In reality, growth of autotrophs and heterotrophs may be limited by various elements, e.g. carbon, nitrogen, phosphorous or iron. In this study we develop a dynamic energy budget model chemostat which has both carbon and nitrogen as currencies, and examine how the dual availability of these elements affects the growth of phytoplankton, trophic transfer to zooplankton, and the resulting stability of the chemostat ecosystem. Both species have two reserve pools to obtain a larger metabolic flexibility with respect to changing external environments. Mineral nitrogen and carbon form the base of the food chain, and they are supplied at a constant rate. In addition, the biota in the chemostat recycle nutrients by means of respiration and excretion, and organic detritus is recycled at a fixed rate. We use numerical bifurcation analysis to assess the model's dynamic behavior. In the model, phytoplankton is nitrogen limited, and nitrogen enrichment can lead to oscillations and multiple stable states. Moreover, we found that recycling has a destabilizing effect on the food chain due to the increased repletion of mineral nutrients. We found that both carbon and nitrogen enrichment stimulate zooplankton growth. Therefore, we conclude that the concept of single-element limitation may not be applicable in an ecosystem context.     

Cascading effects of the flathead grey mullet Mugil cephalus in freshwater eutrophic microcosmos

Flathead grey mullet (Mugil cephalus L.) stocked in fish ponds were long considered to feed primarily on detritus. However, recent research has found that they obtain much of their food from plankton and that they have a detrimental effect on pond zooplankton and large phytoplankton, whilst enhancing small phytoplankton. It has been suggested that flathead grey mullet may also increase the internal phosphorus loading of the ecosystem, which would also increase phytoplankton density. To test whether zooplankton removal or nutrient release from the sediment is the better explanation for phytoplankton enhancement in the presence of flathead grey mullet, the ecosystems of fish-less tanks, tanks with a 60 m mesh filter and tanks stocked at a fish density of 243 g m^-3 were compared. In the presence of flathead grey mullets, cladocerans, ostracods and chironomid larvae became scarcer than in the control tanks, while there were more small phytoplankton and mud snails. The green algae Cladophora sp. did not occur at all. The presence of a mechanical filter also reduced cladoceran, ostracod and chironomid densities and increased phtyoplankton and mud snail density. However, the situation observed in filter tanks was intermediate between that observed in the fish tanks and the control tanks, due to the lower filtering efficiency of the mechanical filter. The organic matter content of the sediment decreased throughout the experiment in the control and filter tanks, but remained stable in fish tanks. Phosphorus and nitrogen concentrations were not affected by any treatment. These results showed that flathead grey mullet enhanced phytoplankton development due to the removal of large cladocerans and not as a consequence of nutrient release. Furthermore, the flathead grey mullet strongly modified the benthic community, probably due to direct predation.     

Coupling of phytoplankton and detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

An oscillating steady state is described of phytoplankton, dominated by Prochlorothrix hollandica and Oscillatoria limnetica, and sestonic detritus in shallow, eutrophic Lake Loosdrecht (The Netherlands). A steady-state model for the coupling of the phytoplankton and detritus is discussed in relation to field and experimental data on phytoplankton growth and decomposition. According to model predictions, the phytoplankton to detritus ratio decreases hyperbolically at increasing phytoplankton growth rate and is independent of a lake's trophic state. The seston in L. Loosdrecht contains more detritus than phytoplankton as will apply to many other lakes. The model provides a basis for estimating the loss rate of the detritus, including decomposition, sedimentation and hydraulic loss. In a shallow lake like L. Loosdrecht detritus will continue to influence the water quality for years.     

Does N2 fixation meet the nitrogen requirements of heterocystous blue-green algae in shallow eutrophic lakes?

Summary Phytoplankton net carbon uptake and nitrogen fixation were studied in two shallow, eutrophic lakes in South Sweden. Ranges of diurnal net carbon uptake were estimated by subtracting 24-h respiration rates corresponding to 5–20% of P _max, respectively, from daytime carbon uptake values. total nitrogen requirement of the phytoplankton assemblage was determined from the diurnal net carbon uptake, assuming a phytoplankton C:N ratio of 9.5:1. Nitrogen supplied by nitrogen fixation only occasionally corresponded to the demands of the total phytoplankton assemblage. When heterocystous algae made up a substantial proportion (10%) of the total phytoplankton biomass, nitrogen fixation could meet the requirements of heterocystous blue-green algae on c. 50% of the sampling occasions. Nitrogen deficiencies in heterocystous algae were most probably balanced by the simultaneous or sequential assimilation of dissolved inorganic nitrogen. It was concluded that uptake of ammonium or nitrate, regenerated from lake seston and sediment, is the main process by which growth of phytoplankton is maintained during summer in the lake ecosystems studied.     

Feeding interactions and foraging of juvenile fish and shrimp in the estuaries of Peter the Great Bay in the summer-fall period

The feeding interrelations and foraging of juvenile fish and shrimp were investigated in estuaries of Peter the Great Bay in summer and fall 2006 and 2007. In total 3483 specimens were analyzed. The trophic spectra of 34 fish species and 3 shrimp species were described for the first time. Three trophic groups were singled out depending on differences in diet compositions: (i) detritophages (3 species of bitterlings—Acanthorhodeus sp., A. chankaensis and Rhodeus sericeus, redlip mullet Liza haematocheila and two species of shrimp-Palaemon macrodactylus and Crangon cf. septemspinosa); (ii) predators (lookup Culter alburnus and Amur catfish Silurus asotus); (iii) euryphages (28 fish species and the shrimp Palaemon paucidens). The detritus food chain was found to be the primary one in estuaries of Peter the Great Bay. Most of the species had two peaks of feeding activity each day. Values of daily feeding rations were determined for 21 fish species; these varied from 1.6% to 8.0% of body weight. Similarity in diet composition was significant only for 10% of the species, which shows a low level of competition for food in the community.     

DECOMPOSITION IN PELAGIC MARINE ECOSYSTEMS

SUMMARY

During the decomposition of plant detritus, complex microbial successions develop which are dominated in the early stages by a number of distinct bacterial morphotypes. The microheterotrophic community rapidly becomes heterogenous and may include cyanobacteria, fungi, yeasts and bactivorous protozoans.

Microheterotrophs in the marine environment may have a biomass comparable to that of all other heterotrophs and their significance as a resource to higher trophic orders, and in the regeneration of nutrients, particularly nitrogen, that support ‘regenerated’ primary production, has aroused both attention and controversy.

Soluble low molecular weight substrates (dissolved organic matter, or DOM) are for the most part rapidly turned over and readily taken up with a high growth efficiency by bacteria although detrital particulate organic material (POM) is turned over slowly and utilized with a low growth efficiency, owing to the structural complexity of the detritus. The presence of appropriate substrate-specific strains of bacteria Is important in the decomposition of both DOM and POM.

Estimates of the transfer of photosynthetically fixed carbon and nitrogen through the pelagic microbial community have recently become widespread. However, the quantification of C and N fluxes through bacteria and microzooplankton is very sensitive to accurate measures of microbial biomass, production, net growth yield, bacterial activity and bactivory by microzooplankton. These processes also vary significantly in both spatial and temporal dimensions during the development and decay of phytoplankton blooms.

Recent attempts to model decomposition processes and C and N fluxes In pelagic marine ecosystems are described. This review examines the most sensitive components and predictions of the models with particular reference to estimates of bacterial production, net growth yield and predictions of N cycling determined by ¹⁵N methodology.     

Nutrient limitation of phytoplankton in a naturally acidic lake (Lake Caviahue, Argentina)

As a result of a low pH, the inorganic carbon of acidic lakes is present as CO₂ at air-equilibrium concentration and is substantially lower than the inorganic carbon concentration in higher-pH waters with bicarbonate. This situation is quite common in artificially acidified lakes and where inorganic carbon is considered the limiting factor in phytoplankton growth. Apart from low inorganic carbon content, Lake Caviahue in Argentina has low nitrogen and high phosphorus content. The aim of this work was to assess the importance of inorganic carbon, phosphorus, and nitrogen, relating data on lake nutrients to phytoplankton species requirements. Lake samples taken in the 2004–2006 period did not show any particular trend in the vertical distribution of the water column of ammonium, inorganic carbon, and phosphorus with reference to either seasonality or depth. A decrease of some 15% in the lake’s phosphorus concentration was observed over the same period. Although the total phytoplankton biomass in Lake Caviahue was similar throughout the period, a seasonal variation was observed. Lab bioassays were carried out with solutions of bicarbonates, ammonium, nitrates, and phosphate. We worked with three species separately, namely, two chlorophytes, Keratococcus rhaphidioides and Watanabea sp.; and one euglenophyte, Euglena mutabilis. Answers to specific nutrient requirements differed for each algal species: both chlorophytes prefer ammonium or nitrates added on their own, whereas the euglenophyte registered a higher growth rate with the joint addition of ammonium and phosphorus. Even when the limiting nutrient(s) for phytoplankton yield and rate varied between species, we observed a tendency for nitrogen limitation in Lake Caviahue.     

Seasonal distribution and changes in the nutritive quality of living,dead and detrital fractions of Zostera marina L.

Samples of eelgrass, Zostera marina L., were collected monthly from December 1974 through December 1975 in a shallow embayment near Beaufort, N.C., and separated into green leaves, dead leaves, and leaf detrital material. Each component was analysed for dry weight, organic matter, inorganic and organic carbon, nitrogen and amino compounds.The standing crop of green and dead blades reached a maximum in April through June, while detrital matter had peaks in December, April and July–September. Inorganic carbon in the three grass fractions showed seasonal variations similar to those observed for epiphytic biomass in previous years, and represented 14, 24, and 30% of the total carbon associated with the green and dead leaves and detrital fragments, respectively. Organic carbon represented a decreasing proportion of the dry weight of these three fractions on a dry weight basis while there was a significant increase in organic carbon on an ash-free dry weight basis in the detrital fragments relative to the head dead blades. During senescence there was a loss of nitrogen from the leaves and an increase in the nitrogen content of the organic matter of the detritus relative to the dead leaves. The latter suggests that there was microbial growth on the detritus and subsequent nitrogen immobilization from the surrounding medium. There were significant decreases of lysine, histidine, arginine, glycine, tyrosine, and glucosamine in the dead leaves relative to the detritus. The glucosamine, derived from N-acetyl-glucosamine, a product of murein which is a component of microbial cell walls, had a seasonal distribution similar to that of the epiphytic community and available inorganic nitrogen in the surrounding water. The relative proportions of N-acetyl-glucosamine, nitrogen and organic carbon were all higher in the fall and winter.     

Influence of nutrient availability on phytoplankton growth and community structure in the Port Adelaide River,Australia: [2pt] bioassay assessment of potential nutrient limitation

We investigated the influence of nutrient availability, specifically nitrogen, phosphorus and silicon on growth and community structure of phytoplankton from the Port Adelaide River estuary, South Australia. Two bioassay experiments were conducted. The first, Nutrich1, involved addition of nutrients in vitro to samples of the natural phytoplankton community from a single location in the upper estuary. The second, Nutrich2, involved nutrient addition and incubation of water from five locations in the estuary following inoculation with a `standardised' phytoplankton assemblage derived from laboratory cultures. In Nutrich1, enrichment with silicon led to greatly enhanced phytoplankton biomass due to increased growth of diatoms. Addition of nitrogen or phosphorus had little effect on phytoplankton growth. In Nutrich2, addition of nitrogen resulted in enhanced growth of phytoplankton in water collected from near the mouth the estuary, but there were no differences in growth among nutrient treatments for the remaining locations. Comparison of phytoplankton growth rate among locations revealed a trend of decreasing growth in moving towards the mouth of the estuary. This trend was unaffected by enrichment with nitrate, phosphate or silicate. We suggest that spatial variation in growth potential within the Port Adelaide River estuary may relate to variation in the concentration of nitrogen as ammonium.     

Phytoplankton exudates provide full nutrition to a subset of accompanying heterotrophic bacteria via carbon,nitrogen and phosphorus allocation

Marine bacteria rely on phytoplankton exudates as carbon sources (DOCp). Yet, it is unclear to what extent phytoplankton exudates also provide nutrients such as phytoplankton-derived N and P (DONp, DOPp). We address these questions by mesocosm exudate addition experiments with spent media from the ubiquitous pico-cyanobacterium Prochlorococcus to bacterial communities in contrasting ecosystems in the Eastern Mediterranean – a coastal and an open-ocean, oligotrophic station with and without on-top additions of inorganic nutrients. Inorganic nutrient addition did not lower the incorporation of exudate DONp, nor did it reduce alkaline phosphatase activity, suggesting that bacterial communities are able to exclusively cover their nitrogen and phosphorus demands with organic forms provided by phytoplankton exudates. Approximately half of the cells in each ecosystem took up detectable amounts of Prochlorococcus-derived C and N, yet based on 16S rRNA sequencing different bacterial genera were responsible for the observed exudate utilization patterns. In the coastal community, several phylotypes of Aureimarina, Psychrosphaera and Glaciecola responded positively to the addition of phytoplankton exudates, whereas phylotypes of Pseudoalteromonas increased and dominated the open-ocean communities. Together, our results strongly indicate that phytoplankton exudates provide coastal and open-ocean bacterial communities with organic carbon, nitrogen and phosphorus, and that phytoplankton exudate serve a full-fledged meal for the accompanying bacterial community in the nutrient-poor eastern Mediterranean.     

Dynamics and limitations of phytoplankton biomass along a gradient in Mwanza Gulf,southern Lake Victoria (Tanzania)

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Do marine phytoplankton follow Bergmann's rule sensu lato?

Global warming has revitalized interest in the relationship between body size and temperature, proposed by Bergmann's rule 150 years ago, one of the oldest manifestations of a ‘biogeography of traits’. We review biogeographic evidence, results from clonal cultures and recent micro‐ and mesocosm experiments with naturally mixed phytoplankton communities regarding the response of phytoplankton body size to temperature, either as a single factor or in combination with other factors such as grazing, nutrient limitation, and ocean acidification. Where possible, we also focus on the comparison between intraspecific size shifts and size shifts resulting from changes in species composition. Taken together, biogeographic evidence, community‐level experiments and single‐species experiments indicate that phytoplankton average cell sizes tend to become smaller in warmer waters, although temperature is not necessarily the proximate environmental factor driving size shifts. Indirect effects via nutrient supply and grazing are important and often dominate. In a substantial proportion of field studies, resource availability is seen as the only factor of relevance. Interspecific size effects are greater than intraspecific effects. Direct temperature effects tend to be exacerbated by indirect ones, if warming leads to intensified nutrient limitation or copepod grazing while ocean acidification tends to counteract the temperature effect on cell size in non‐calcifying phytoplankton. We discuss the implications of the temperature‐related size trends in a global‐warming context, based on known functional traits associated with phytoplankton size. These are a higher affinity for nutrients of smaller cells, highest maximal growth rates of moderately small phytoplankton (ca. 10² µm³), size‐related sensitivities for different types of grazers, and impacts on sinking rates. For a phytoplankton community increasingly dominated by smaller algae we predict that: (i) a higher proportion of primary production will be respired within the microbial food web; (ii) a smaller share of primary production will be channeled to the classic phytoplankton – crustacean zooplankton – fish food chain, thus leading to decreased ecological efficiency from a fish‐production point of view; (iii) a smaller share of primary production will be exported through sedimentation, thus leading to decreased efficiency of the biological carbon pump.     

Nutrient leaching and settling rate characteristics of the faeces of Atlantic salmon (Salmo salar L.) and the implications for modelling of solid waste dispersion

The present study determined the settling velocity and leaching rates of carbon and nitrogen from the faecal matter of Atlantic salmon (Salmo salar L.) fed two commercial diets, high energy (HE) and standard, in three separate experiments (December, March and May) which coincided with winter, spring and summer. Faecal settling velocities for individual samples were in the range of 3.7–9.2 cm s^?1. There were significant differences (P < 0.05) in settling velocities among the three experiments, with lowest velocities measured in faecal samples collected in winter and highest velocities measured in samples collected in summer. There were no significant differences (P > 0.05) in faecal settling velocity among fish fed the different diet types. Faecal nutrient content in the different treatment groups prior to leaching was 269–318 mg g^?1 for carbon and 28–37 mg g^?1 for nitrogen. Both carbon and nitrogen contents were higher in faeces from fish fed the standard diet than in fish fed the HE diet (P < 0.05). Thus the use of HE diets resulted in a 12% reduction in faecal carbon content and an 8% reduction in faecal nitrogen when compared with standard diets. There were no consistent differences in faecal carbon among samples collected during the three experiments; however, significant differences in faecal nitrogen content were detected in samples collected on the three different occasions. Leaching of faecal carbon and faecal nitrogen ranged between 4–14% and 9–16% of the original amount, respectively, after 2.5‐min immersion in sea water, although there was no further significant (P > 0.05) leaching after this time. No significant difference in nutrient leaching rate was found in faeces of fish fed HE and standard diets, and no significant differences in leaching rates were apparent among samples collected at different times of the year. These values suggest there may be an overestimation by dispersion models of the amount of nutrients entering seabed sediments. The amount of nutrients leaching from faeces may also have an important role in nutrient flux in the water column.     

武汉东湖颗粒悬浮物的结构与元素组成

本研究于1989-1990对武汉东湖营养水平不同的二个湖区的颗粒悬浮物的干物质结构和元素组成进行了分析。综合平均值表明,浮游动物的现存量约为浮游植物的1/4,浮游动物群落以小型的原生动物和轮虫占优势。从年平均值来看,浮游生物的干重占颗粒悬浮物干物质的2.5-7.6%,浮游生物碳量占颗粒悬浮物碳量的4.0-9.8%;颗粒悬浮物的碳/氮比与一般浮游植物的比值相似,但明显大于多数浮游动物;颗粒悬浮物的碳与干物重之比约为一般浮游生物的3/4;颗粒悬浮物的灰分含量约为45%,显着高于除硅藻以外的其它浮游生物。从数量上来看,有机碎屑是东湖生态系统颗粒悬浮物最重要的组成部分,而活体浮游生物只占颗粒悬浮物很小的一部分(<10%);这意味着在东湖来自以浮游植物为核心的食物网的有机碎屑的形成速率显着大于有机碎屑的矿化速率。       

Allocation to reproduction in the chaparral shrub,Diplacus aurantiacus

Summary The semi-drought-deciduous shrub, Diplacus aurantiacus, allocates a large, relatively constant proportion of carbon and nitrogen to sexual reproduction. Experimental manipulation at a site in the chaparral of coastal central California showed that both reproduction and vegetative growth were strongly limited by water and little affected by shade or by addition of nutrients unless accompanied by water. Potential competition for carbon between growth and reproduction is reduced by photosynthesis within reproductive structures; competition is also constrained by localization of translocation. Results are discussed in relation to the hypothesis that allocation to reproduction in Diplacus has been selected to maximize reproductive success.     

Nutrient removal from sewage by an artificial food web system composed of phytoplankton and Daphnia magna

In order to remove nutrients from sewage, ecotechnology with an artificial food web composed of phytoplankton and Daphnia magna was used. To optimise performance of the system, phytoplankton growth, zooplankton growth, and a continuous-flow system were used. For phytoplankton growth, stirring was 6.7 times faster than the settling in growth rate of Scenedesmus. Zooplankton growth was not influenced by phytoplankton succession, and the specific production coefficient of D. magna was 110.4 mg Daphnia dry weight (DW) per mg chlorophyll a (Chl a). Results indicated that removal of nutrients was better in a long hydraulic residence time (HRT) system than in a system with short HRT. The optimum retention time was found to be 3 days for the phytoplankton chamber and 1.5 days for the subsequent D. magna chamber, respectively, with total retention time of the combined chambers being kept at 4.5 days. When a pilot plant was operated under these conditions, the removal rates of total nitrogen (TN) and total phosphorus (TP) were 68 and 56%, respectively. In the material budget of TN, 32% of inputs passed on to effluent, 39% to sludge, 27% to air and 2% to harvested Daphnia. For TP, 44% of inputs passed on to effluent, 51% to sludge and 4% to Daphnia.     

The effect of increased nitrogen load on phytoplankton in a phosphorus‐limited lake

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Unimodal size scaling of phytoplankton growth and the size dependence of nutrient uptake and use

Phytoplankton size structure is key for the ecology and biogeochemistry of pelagic ecosystems, but the relationship between cell size and maximum growth rate (μ_max) is not yet well understood. We used cultures of 22 species of marine phytoplankton from five phyla, ranging from 0.1 to 10⁶ μm³ in cell volume (V_cell), to determine experimentally the size dependence of growth, metabolic rate, elemental stoichiometry and nutrient uptake. We show that both μ_max and carbon‐specific photosynthesis peak at intermediate cell sizes. Maximum nitrogen uptake rate (V_maxN) scales isometrically with V_cell, whereas nitrogen minimum quota scales as V_cell^0.84. Large cells thus possess high ability to take up nitrogen, relative to their requirements, and large storage capacity, but their growth is limited by the conversion of nutrients into biomass. Small species show similar volume‐specific V_maxN compared to their larger counterparts, but have higher nitrogen requirements. We suggest that the unimodal size scaling of phytoplankton growth arises from taxon‐independent, size‐related constraints in nutrient uptake, requirement and assimilation.     

Nutrient limitation in Crater Lake,Oregon

Experiments were carried out to determine what nutrient (or nutrients) was primarily responsible for limiting phytoplankton productivity in ultraoligotrophic Crater Lake. The experiments included in situ and laboratory nutrient addition bioassays utilizing the natural phytoplankton community, Selenastrum capricornutum bottle assays, photosynthetic responses, photosynthetic carbon metabolism, and response of dark uptake of ¹⁴CO₂ with the addition of NH ₄ ⁺ . The results suggested that a trace metal(s) or its availability was the primary factor limiting the epilimnetic phytoplankton productivity. Nitrogen was extremely low, and quickly became limiting with the addition of trace metals and a chelator. Iron is the most likely candidate as the limiting nutrient. Trace metals and nitrogen are also both important in limiting phytoplankton at 100 m, a depth where biologically mediated turnover of nutrients seems to be more important.     

Wood Decomposition Following a Perennial Lupine Die-Off: A 3-Year Litterbag Study

Woody debris is a conspicuous feature of many ecosystems and can be a large pool of stored carbon and nutrients. In the California coastal prairie, yellow bush lupines (Lupinus arboreus) experience mass die-offs, producing large quantities of woody detritus. Live lupines are fed upon by the stem-boring caterpillars of the ghost moth, Hepialus californicus, and outbreaks of ghost moths are one factor contributing to lupine die-offs. A common detritivore, the terrestrial isopod Porcellio scaber, frequently inhabits ghost moth tunnels in lupine wood. We used a litterbag experiment to test the hypothesis that H. californicus increases decomposition of woody lupine detritus by facilitating its use by P. scaber. Isopod access to wood was crossed with simulated ghost moth boring to measure the independent and interactive effects of these two arthropods on total mass loss, as well as on carbon, nitrogen, and lignin dynamics. Isopods initially colonized litterbags but were not more abundant on L. arboreus logs that had simulated ghost moth boring than on logs without boring. They were rare in litterbags collected at 12 months or later and had no effect on wood decomposition. Simulated ghost moth boring increased wood decomposition (P = 0.0021), from 50.5 to 55.1% mass loss after 3 years. This effect was likely due to increased surface area for microbial utilization of the wood. Lupine wood had an initial lignin content of 14.70 ± 0.67%, but lignin did not appear to decompose during the 3 years of this study, and by the end of the experiment accounted for 32.6 ± 1.12% of the remaining wood. Neither ghost moth boring nor isopod access affected lignin loss. Lupine wood from a die-off in 2002 was estimated to have contained three times more nitrogen per unit area than the yearly input of annual grass litter. The slow decomposition of lupine wood, however, restricts the rate at which nitrogen is released into the soil and results in the storage of carbon and nutrients in lupine wood for several years following such die-offs.