Elemental Composition Of Phytoplankton Research Articles

A model of time-dependent macromolecular and elemental composition of phytoplankton

Phytoplankton Chl:C:N:P ratios are important from both an ecological and a biogeochemical perspective. We show that these elemental ratios can be represented by a phytoplankton physiological model of low complexity that includes major cellular macromolecular pools. In particular, our model resolves time-dependent intracellular pools of chlorophyll, proteins, nucleic acids, carbohydrates/lipids, and N and P storage. Batch culture data for two diatom and two prasinophyte species are used to constrain parameters that represent specific allocation traits and strategies. A key novelty is the simultaneous estimation of physiological parameters for two phytoplankton groups of such different sizes. The number of free parameters is reduced by assuming (i) allometric scaling for maximum uptake rates, (ii) shared half-saturation constants for synthesis of functional macromolecules, (iii) shared exudation rates of functional macromolecules across the species. The rationale behind this assumption is that across the different species, the same or similar processes, enzymes, and metabolites play a role in key physiological processes. For the turnover numbers of macromolecular synthesis and storage exudation rates, differences between diatoms and prasinophytes need to be taken into account to obtain a good fit. Our model fits suggest that the parameters related to storage dynamics dominate the differences in the C:N:P ratios between the different phytoplankton groups. Since descriptions of storage dynamics are still incomplete and imprecise, predictions of C:N:P ratios by phytoplankton models likely have a large uncertainty.

Effect of Flocks of Anseriform Birds on Seston and Phytoplankton in Lakes of the Taimyr Peninsula

The effect of molting anseriform birds on the structure and elemental composition of phytoplankton (seston) has been assessed in 20 Arctic lakes of the Taimyr Peninsula. In lakes (part of the lake) inhabited by ~50–700 birds of six species, the average stoichiometric ratio N : P (mol : mol) was statistically significantly lower than in lakes without anseriforms: 15.8 ± 1.4 and 22.4 ± 2.7, respectively. There was also a tendency of higher average specific electrical conductivity in the lakes with the birds, 113 ± 32 µS/cm, when compared with those without anseriforms, 60 ± 18 µS/cm. The differences could be explained with high probability by the effect of guanotrophication, namely, by a flow in water of metabolites of molting anseriforms. The total biomass of phytoplankton and proportions of algal taxa and cyanobacteria in the total biomass did not differ statistically significantly in lakes with and without molting anseriforms. Therefore, under guanotrophication, the main threat of eutrophication was absent: an increase of biomass of cyanobacteria, causing the nuisance “bloom” of water. Moreover, an opposite tendency occured: in lakes with molting anseriforms, the proportion of cyanobacteria in total biomass of phytoplankton was on average lower than that in lakes without the birds, 16.2 ± 5.3% and 30.8 ± 9.3%, respectively. Thus, a hypothesis was confirmed that artificial guanotrophication should be regarded as a suitable ecotechnology for the increase of productivity of oligotrophic Arctic lakes.

Ecological response of Rotaria rotatoria (Bdelloid Rotifera) to unbalanced nitrogen in food: experimental insights from life history strategy and feeding behavior

ABSTRACT Nitrogen (N) cycle in ecosystems has been overbalanced by human activities. However, it remains uncertain whether the altered N supply results in a change in the elemental composition of phytoplankton and, consequently, affects the life history strategy of zooplankton. To investigate these impacts, a simple lab-based food chain was established. Results show that lack or excess of nitrogen reduced algal density, cell volume, growth rate and chlorophyll content. Moreover, N content in algae significantly increased with increased N concentration in the medium, and reached saturation at concentrations ≥5 mg·L−1. Feeding on algae grown in a low-nitrogen or no nitrogen mediums resulted in faster decline in age-specific survival of rotifers, and slower population growth, as well as longer generation time. In order to make up for nutritional shortage, grazing and filtration rates increased. On the other hand, rotifers feeding on algae grown in high-N mediums (A50 and A200) had significantly shorter average lifespan and life expectancy at hatching. Therefore, nitrogen imbalances have adverse effects on the growth, development and reproduction of both primary producers and herbivorous zooplankton in the food chain.

A meta-analysis on environmental drivers of marine phytoplankton C : N : P

Abstract. The elemental stoichiometry of marine phytoplankton plays a critical role in global biogeochemical cycles through its impact on nutrient cycling, secondary production, and carbon export. Although extensive laboratory experiments have been carried out over the years to assess the influence of different environmental drivers on the elemental composition of phytoplankton, a comprehensive quantitative assessment of the processes is still lacking. Here, we synthesized the responses of P:C and N:C ratios of marine phytoplankton to five major drivers (inorganic phosphorus, inorganic nitrogen, inorganic iron, irradiance, and temperature) by a meta-analysis of experimental data across 366 experiments from 104 journal articles. Our results show that the response of these ratios to changes in macronutrients is consistent across all the studies, where the increase in nutrient availability is positively related to changes in P:C and N:C ratios. We found that eukaryotic phytoplankton are more sensitive to the changes in macronutrients compared to prokaryotes, possibly due to their larger cell size and their abilities to regulate their gene expression patterns quickly. The effect of irradiance was significant and constant across all studies, where an increase in irradiance decreased both P:C and N:C. The P:C ratio decreased significantly with warming, but the response to temperature changes was mixed depending on the culture growth mode and the growth phase at the time of harvest. Along with other oceanographic conditions of the subtropical gyres (e.g., low macronutrient availability), the elevated temperature may explain why P:C is consistently low in subtropical oceans. Iron addition did not systematically change either P:C or N:C. Overall, our findings highlight the high stoichiometric plasticity of eukaryotes and the importance of macronutrients in determining P:C and N:C ratios, which both provide us insights on how to understand and model plankton diversity and productivity.

The effects of absolute and relative nutrient concentrations (N/P) on phytoplankton in a subtropical reservoir

Abstract The elemental composition of phytoplankton is a critical factor for primary production and nutrient recycling. The increase anthropogenic nutrient input into freshwater ecosystems is affecting phytoplankton assemblage structure and its stoichiometry. Reservoirs of South China generally show low level of phosphate and it is not clear how phytoplankton can grow and occasionally bloom in such conditions. Therefore, an indoor experiment was conducted to investigate the response of natural phytoplankton communities to 25 levels of supplied nitrogen to phosphorus ratios (N/P), arising from the combination of 5 levels of N and P. Our aim was to check the effects of absolute and relative N and P on phytoplankton growth and structure. We assumed that Alkaline Phosphatase (APA) provides a way to use alternative P resource. Our hypotheses include: (1) phytoplankton stoichiometry would be in homeostasis (sensu Sterner & Elser, 2002) under different N and P treatments; (2) absolute nutrient values rather than its ratio matters for phytoplankton assemblage; and (3) phytoplankton cell size declines to lower P requirements facing P limitation. Results showed that phytoplankton in this subtropical reservoir use alternative P sources via APA to support its growth. The absolute values of N and P instead and not their ratios were important for phytoplankton. The N/P ratio cannot be a reliable indicator for nutrient limitation or phytoplankton shift. During the fast-growing phase, their elemental contents were not sensitive to supplied nutrients, and their stoichiometry was more constrained compared to that in slow-growing phase. The plasticity of cellular stoichiometry was mainly due to the variations of cellular P. Phytoplankton stoichiometry was weakly homeostatic, and this mechanism provides a strategy to keep growth relatively stable in a variable nutrient environment.

The Temperature Dependence of Phytoplankton Stoichiometry: Investigating the Roles of Species Sorting and Local Adaptation.

The elemental composition of phytoplankton (C:N:P stoichiometry) is a critical factor regulating nutrient cycling, primary production and energy transfer through planktonic food webs. Our understanding of the multiple direct and indirect mechanisms through which temperature controls phytoplankton stoichiometry is however incomplete, increasing uncertainty in the impacts of global warming on the biogeochemical functioning of aquatic ecosystems. Here, we use a decade-long warming experiment in outdoor freshwater ponds to investigate how temperature-driven turnover in species composition and shifts in stoichiometric traits within species through local thermal adaptation contribute to the effects of warming on seston stoichiometry. We found that experimental warming increased seston C:P and N:P ratios, while the C:N ratio was unaffected by warming. Temperature was also the dominant driver of seasonal variation in seston stoichiometry, correlating positively with both C:P and N:P ratios. The taxonomic composition of the phytoplankton community differed substantially between the warmed and ambient treatments indicating that warming resulted in differential sorting of species from the regional pool. Furthermore, taxonomic composition also changed markedly over the year within each of the warmed and ambient treatments, highlighting substantial temporal turnover in species. To investigate whether local adaptation also played an important role in shaping the effects of warming on seston stoichiometry, we isolated multiple strains of the cosmopolitan alga, Chlamydomonas reinhardtii from across the warmed and ambient mesocosms. We found that warmed isolates had higher C:P and N:P ratios, shifts that were comparable in direction and magnitude to the effects of warming on seston stoichiometry. Our results suggest that both species sorting and local adaptation are likely to play important roles in shaping the effects of warming on bulk phytoplankton stoichiometry and indicate that major shifts in aquatic biogeochemistry should be expected in a warmer world.

Interactions between Thermal Acclimation, Growth Rate, and Phylogeny Influence Prochlorococcus Elemental Stoichiometry.

Variability in plankton elemental requirements can be important for global ocean biogeochemistry but we currently have a limited understanding of how ocean temperature influences the plankton C/N/P ratio. Multiple studies have put forward a ‘translation-compensation’ hypothesis to describe the positive relationship between temperature and plankton N/P or C/P as cells should have lower demand for P-rich ribosomes and associated depressed QP when growing at higher temperature. However, temperature affects many cellular processes beyond translation with unknown outcomes on cellular elemental composition. In addition, the impact of temperature on growth and elemental composition of phytoplankton is likely modulated by the life history and growth rate of the organism. To test the direct and indirect (via growth rate changes) effect of temperature, we here analyzed the elemental composition and ratios in six strains affiliated with the globally abundant marine Cyanobacteria Prochlorococcus. We found that temperature had a significant positive effect on the carbon and nitrogen cell quota, whereas no clear trend was observed for the phosphorus cell quota. The effect on N/P and C/P were marginally significantly positive across Prochlorococcus. The elemental composition and ratios of individual strains were also affected but we found complex interactions between the strain identity, temperature, and growth rate in controlling the individual elemental ratios in Prochlorococcus and no common trends emerged. Thus, the observations presented here does not support the ‘translation-compensation’ theory and instead suggest unique cellular elemental effects as a result of rising temperature among closely related phytoplankton lineages. Thus, the biodiversity context should be considered when predicting future elemental ratios and how cycles of carbon, nitrogen, and phosphorus may change in a future ocean.

Food quality dominates the impact of food quantity on Daphnia life history: possible implications for re-oligotrophication

The elemental composition of phytoplankton is highly variable compared to the relatively narrow stoichiometry of zooplankton grazers. Using a full factorial design, we tested the effects of alterations in algal elemental composition (i.e., food quality) combined with food quantity on the life history of a Daphnia galeata clone from Lake IJsselmeer. Lower food quality reduced survival, growth, and reproduction. Food quantity became important at high food quality only. The strong effect of food quality indicates the potential for a stoichiometric bottleneck in Lake IJsselmeer, resulting in less high quality food for higher trophic levels as a result of re-oligotrophication.

Strong latitudinal patterns in the elemental ratios of marine plankton and organic matter

The elemental composition of marine organic matter is used to infer a variety of oceanic ecosystem processes. A compilation of observational data suggests that elemental ratios differ substantially from the Redfield ratio, but exhibit a clear latitudinal trend. Nearly 75 years ago, Alfred C. Redfield observed a similarity between the elemental composition of marine plankton in the surface ocean and dissolved nutrients in the ocean interior1. This stoichiometry, referred to as the Redfield ratio, continues to be a central tenet in ocean biogeochemistry, and is used to infer a variety of ecosystem processes, such as phytoplankton productivity and rates of nitrogen fixation and loss2,3,4. Model, field and laboratory studies have shown that different mechanisms can explain both constant and variable ratios of carbon to nitrogen and phosphorus among ocean plankton communities. The range of C/N/P ratios in the ocean, and their predictability, are the subject of much active research5,6,7,8,9,10,11,12. Here we assess global patterns in the elemental composition of phytoplankton and particulate organic matter in the upper ocean, using published and unpublished observations of particulate phosphorus, nitrogen and carbon from a broad latitudinal range, supplemented with elemental data for surface plankton populations. We show that the elemental ratios of marine organic matter exhibit large spatial variations, with a global average that differs substantially from the canonical Redfield ratio. However, elemental ratios exhibit a clear latitudinal trend. Specifically, we observed a ratio of 195:28:1 in the warm nutrient-depleted low-latitude gyres, 137:18:1 in warm, nutrient-rich upwelling zones, and 78:13:1 in cold, nutrient-rich high-latitude regions. We suggest that the coupling between oceanic carbon, nitrogen and phosphorus cycles may vary systematically by ecosystem.

The unique biogeochemical signature of the marine diazotroph trichodesmium.

The elemental composition of phytoplankton can depart from canonical Redfield values under conditions of nutrient limitation or production (e.g., N fixation). Similarly, the trace metal metallome of phytoplankton may be expected to vary as a function of both ambient nutrient concentrations and the biochemical processes of the cell. Diazotrophs such as the colonial cyanobacteria Trichodesmium are likely to have unique metal signatures due to their cell physiology. We present metal (Fe, V, Zn, Ni, Mo, Mn, Cu, Cd) quotas for Trichodesmium collected from the Sargasso Sea which highlight the unique metallome of this organism. The element concentrations of bulk colonies and trichomes sections were analyzed by ICP-MS and synchrotron x-ray fluorescence, respectively. The cells were characterized by low P contents but enrichment in V, Fe, Mo, Ni, and Zn in comparison to other phytoplankton. Vanadium was the most abundant metal in Trichodesmium, and the V quota was up to fourfold higher than the corresponding Fe quota. The stoichiometry of 600C:101N:1P (mol mol−1) reflects P-limiting conditions. Iron and V were enriched in contiguous cells of 10 and 50% of Trichodesmium trichomes, respectively. The distribution of Ni differed from other elements, with the highest concentration in the transverse walls between attached cells. We hypothesize that the enrichments of V, Fe, Mo, and Ni are linked to the biochemical requirements for N fixation either directly through enrichment in the N-fixing enzyme nitrogenase or indirectly by the expression of enzymes responsible for the removal of reactive oxygen species. Unintentional uptake of V via P pathways may also be occurring. Overall, the cellular content of trace metals and macronutrients differs significantly from the (extended) Redfield ratio. The Trichodesmium metallome is an example of how physiology and environmental conditions can cause significant deviations from the idealized stoichiometry.

Evolutionary inheritance of elemental stoichiometry in phytoplankton

The elemental composition of phytoplankton is a fusion of the evolutionary history of the host and plastid, resulting in differences in genetic constraints and selection pressures associated with environmental conditions. The evolutionary inheritance hypothesis predicts similarities in elemental composition within related taxonomic lineages of phytoplankton. To test this hypothesis, we measured the elemental composition (C, N, P, S, K, Mg, Ca, Sr, Fe, Mn, Zn, Cu, Co, Cd and Mo) of 14 phytoplankton species and combined these with published data from 15 more species from both marine and freshwater environments grown under nutrient-replete conditions. The largest differences in the elemental profiles of the species distinguish between the prokaryotic Cyanophyta and primary endosymbiotic events that resulted in the green and red plastid lineages. Smaller differences in trace element stoichiometry within the red and green plastid lineages are consistent with changes in trace elemental stoichiometry owing to the processes associated with secondary endosymbioses and inheritance by descent with modification.

Exhaustion of nitrate terminates a phytoplankton bloom in Funka Bay, Japan:change in SiO4:NO3 consumption rate during the bloom

Time series observations over 36 h were carried out in Funka Bay, Japan, to complement a 6 yr nutrient-dynamics study during a diatom spring bloom, especially focusing on nitrate and silicate as limiting nutrients. At the beginning of the study, nutrient concentrations were high (NO 3 = 10 μM, SiO 4 = 20 μM and PO 4 = 0.8 μM). Nitrate was depleted first during the night while silicate and phosphate remained at 10 and 0.3 μM, respectively. High nitrate reductase activity was observed the following morning, indicating the assimilation of nitrate in situ by the dominant phytoplankter, Chaetoceros socialis. The consumption ratio of SiO 4 :NO 3 increased by 3 times from 0.83 before the peak to 2.5 after the peak of the bloom. This change was also obvious in the elemental composition of phytoplankton, showing an increase in Si:C and Si:N ratios by a factor of 2 to 4 after the peak of the bloom, while C:N ratio remained constant at around 7. As a consequence of this change in the consumption rate of SiO 4 and NO 3 , both silicate and nitrate were finally depleted in the photic zone.

The Role of Episodic Atmospheric Nutrient Inputs in the Chemical and Biological Dynamics of Oceanic Ecosystems

ATMOSPHERIC INPUTS of iron and nitrogen have been hypothesized to play an important role in the chemical and biological dynamics of openocean ecosystems (Menzel and Spaeth, 1962; Duce, 1986). This hypothesis has stimulated considerable discussion and controversy (see Martin, 1991, this issue; Miller et al., 1991, this issue; Morel et al., 1991, this issue). Much of this discussion has focused on the relative importance of atmospheric nutrient input rates over large time and space scales, the physiology and chemistry of iron nutrition, the elemental composition of phytoplankton (e.g., Fe/C), and the interpretation of assays for iron limitation ofphytoplankton growth at a particular time and place. An interdisciplinary analysis has led to the realization that biological responses of oceanic ecosystems are likely to be dependent not only on biological requirements and the large-scale variability in annual-average atmospheric inputs but also on the temporal and spatial variability of these inputs, on the chemical and biological processes controlling the fate of those inputs, and on the biological, chemical, and physical interactions controlling the dynamics of open-ocean ecosystems receiving those inputs. The results of this interdisciplinary analysis are summarized below.