Urea uptake by phytoplankton at various stages of nutrient depletion

Uptake of ¹⁴C-urea by Thalassiosira pseudonana and Skeletonemacostatum grown in batch culture with NO₂^– and NO₃^–as nitrogen sources was measured under three conditions: pre-depletion(when nitrogenous nutrient was present in the culture mediumat saturating concentrations), at-depletion (when nitrogenousnutrient could no longer be detected), and several hours post-depletion.V_max-urea, the initial instantaneous uptake rate, remained constantunder all three conditions, and was in excess of uptake ratesrequired for cellular doubling. Variations in uptake under thethree conditions were observed, as functions of the length oftime over which uptake was observed and the growth rate of theculture. The maximum instantaneous uptake rate was not differentfor the three conditions; variations in uptake were due to theperiod of time over which the maximum uptake rate was maintained.The ability of cells to take up urea rapidly, even when adequatelynourished by NO₂^– and NO₃^–, could be of significancein a low and variable urea-nutrient regime in the natural environment.     

Scaling-up from nutrient physiology to the size-structure of phytoplankton communities

In many community assemblages, the abundance of organisms isa power-law function of organism size. In phytoplankton communities,changes in size structure associated with increases in resourceavailability and total biomass have often been interpreted asa release from grazer control. A metapopulation-like approachis used to scale up from the individual physiological responsesto environmental conditions to community size structure assumingthe community taxonomic composition reflects the species pool.We show that the size scaling of cellular nutrient requirementsand growth can cause (1) the power-law relationship betweencell size and abundance, (2) dominance of small phytoplanktoncells under oligotrophic conditions, and (3) relative increasein abundance of larger phytoplankton cells under eutrophic conditions.If physiological differences associated with the taxonomic compositionof different community size fractions are considered, then themodel can replicate detailed field observations such as theabsence of small, slow-growing Prochlorococcus spp. and therelative dominance of large diatom species in nutrient-rich,upwelling regions of the ocean. This paper was presented in a session on "Size Structure ofPlankton Communities", at the ASLO Summer International Meeting,held in Santiago de Compostela, Spain, between 19 and 24 June,and coordinated by Xabier Irigoien, Roger Harris and Angel Lopez-Urrutia.     

Vertical nutrient and phytoplankton distribution in relation to physical stability

The vertical distribution of nutrients and phytoplankton in relation to water stability in Saronicos Gulf, Greece, was examined during mixing and during water stratification. Phosphate, nitrate and phytoplankton were stratified during mixing (February), and phytoplankton was well stratified, mainly in April. Thus nutrient and phytoplankton vertical distribution do not always follow the motion of the water and eutrophic conditions favour nutrient and phytoplankton stratification.     

Quantifying the impact of periphytic algae on nutrient availability for phytoplankton

SUMMARY.

• 1 Recent laboratory studies demonstrate that periphytic algae growing on the sediment surface reduce nutrient availability in the overlying water. Consequently, periphytic algae may competitively reduce growth of phytoplankton.
• 2 The aim of this study was to quantify the competitive impact of sediment-attached periphytic algae on phytoplankton in the presence of all other factors simultaneously affecting nutrient dynamics in natural systems.
• 3 In enclosure experiments, performed in three lakes of different productivity, the periphytic algal biomass was manipulated. When compared to enclosures with high biomass of periphytic algae, those with reduced biomass showed an increase in total phosphorus concentration in the water of 32–44%. Extrapolation of the experimental results to whole lakes predicts an increase in original total phosphorus concentration of between 1.5% and 8.0%. According to existing regressions between total phosphorus and phytoplankton chlorophyll, the potential increase in original phytoplankton biomass will be between 2.5% and 12.6%.
• 4 With respect to the shallow parts of lakes, my results support the conclusions revealed from laboratory studies that periphytic algae have a significant impact on the phosphorus concentration in the overlying water. However, when considering whole-lake dynamics, the competitive impact of periphytic algae on phytoplankton biomass development is probably of minor importance.
• 5 Rather, the main competitive advantage of growing on the sediment surface, compared to in the water, may be the exclusive access to nutrients in the sediment.

     

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.     

Size-fractionated productivity and nutrient dynamics of phytoplankton in subtropical coastal environments

It is now well established that the size distribution of phytoplankton plays an important role in primary production processes and nutrient dynamics of coastal environment. In situ observations showed that nanophytoplankton (3–20 μm) contributed 72.08% and58.18% of phytoplankton biomass and 58.32% and 41.14% of primary productivity to Xiamen Western Waters and the northern Taiwan Strait, respectively; picophytoplankton (0.2–3 μm) dominated the biomass (64.70%) and productivity (66.09%) in the southern Taiwan Strait. Furthermore, nanophytoplankton accounted for 75% of phosphate uptake with the highest rate constant (8.3×10^-5 s^-1) and uptake rate in unit water volume (5.4×10^-5 mmol dm^-3s^-1); picophytoplankton had the highest uptake rate in unit biomass (5.4×10^-5 mmol mg^-1s^-1) and photosynthetic index (3.8 mgC mgChl a^-1h^-1). All the results highlighted the remarkable characteristics of small size ranged (0.2–20 μm) phytoplankton in subtropical coastal environments: main contributor to phytoplankton biomass and production, high efficiency on organic carbon production and nutrient recycling. The far reaching environmental and ecological implications were discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Physiological indicators of nutrient deficiency in phytoplankton in southern Chilean lakes

We assessed the nutrient status of phytoplankton in 28 lakes in southern Chile using two types of physiological indicators: specific alkaline phosphatase activity, and the elemental composition (carbon, nitrogen, and phosphorus) of seston. Alkaline phosphatase activity ranged from 0.001 to 0.11 mol P g chl^–1 h^–1, with P-deficiency indicated in about one-half the study lakes. C:N ranged from 3.9 to 24, C:P ranged from 86 to 919, and N:P ranged from 8.7 to 99. C:P and N:P ratios greater than the Redfield ratio were common, suggesting P deficiency in many of the lakes. C:N ratios were not generally indicative of N deficiency. Previous studies have suggested N may be the primary limiting nutrient in southern Chilean lakes, but our results indicate that P should not be discounted as a limiting nutrient.     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Relationship between nutrient concentration,phytoplankton density,and zooplankton density in nutrient enriched experimental ponds

The effects of different levels of nutrient input on the plankton community was investigated in a two-year controlled fertilization study of eight experimental ponds. There were four treatments, each replicated: a control, to which no fertilizer was added, and three levels of nutrient addition. Limnological parameters including phytoplankton and zooplankton densities were measured frequently during both summers and less frequently during the rest of the year. Inorganic nitrogen and phosphorus concentrations in the treated ponds increased. Phytoplankton and zooplankton density increased with treatment level but was variable. There was a limited relationship between the average chlorophyll a concentration per summer and the average cladoceran dry weight per summer. Above chlorophyll a concentrations greater than 60–70 mg/m³ other factors such as a pH zooplankton mortality effect, prevailed.     

Effects of nutrient and light limitation on the biochemical composition of phytoplankton

Three marine phytoplankters (Isochrysis galbana, Chaetoceros calcitrans andThalassiosira pseudonana), commonly used in the culture of bivalve larvae, were grown in batch or semi-continuous cultures. Changes in protein, carbohydrate, lipid and some fatty acids were measured as growth became limited by nitrogen, silicon, phosphorus or light. Under N starvation (2 d) the % lipid remained relatively constant, while% carbohydrate increased and% protein decreased in all 3 species compared to cells growing under no nutrient limitation. Under Si starvation (6 h) there was no change in lipid, protein or carbohydrates. The amount of two fatty acids, 20 : 53 and 22 : 63 remained relatively constant under N, P and Si starvation, exept for a sharp drop in the cells of P-starvedT. pseudonana. However, there were pronounced species differences withI. galbana containing significantly less 20 : 5 3 thanC. calcitrans orT. pseudonana. Under light limitation the amount of lipid per cell showed no consistent trend over a range of irradiances for all 3 species. The amount of N per cell (an index of protein content) as a function of irradiance, was relatively constant forI. galbana andT. pseudonana, while the amount of N per cell was lower under low irradiances forC. calcitrans. These examples of changes in protein, carbohydrate, lipid and certain fatty acids under nutrient (N, Si or P) or light limitation, emphasize the importance of knowing the phase (e.g. logarithmic vs stationary) of the growth curve in batch cultures, since the nutritional value of the phytoplankters could change as cultures become dense and growth is terminated due to nutrient or light limitation.Presented at the XIIIth International Seaweed Symposium, University of British Columbia, Vancouver, Canada, August 1989.     

On limiting nutrient patchiness and phytoplankton growth: a conceptual approach

A theoretical framework is developed to explore the effectsof limiting nutrient patchiness on phytoplankton growth. Growthrate is represented as a function of the average ambient substrateconcentration in the medium, the degree of patchiness and thepatch duration. Phytoplankton growth, in relation to the externalsubstrate concentration, is mediated by the cell quota for thelimiting nutrient. Two general conclusions can be drawn from this study. Firstthe degree of patchiness in the environment can affect individualgrowth rates and thus alter community structure even thoughthere is no change in the average ambient nutrient concentration.Second, for patch-adapted populations, the apparent K_s for growthcan be lowered significantly by making the distribution of thelimiting nutrient patchy with respect to time. The insightswhich this model provides into future experimental methodologiesare also discussed. ¹ Address for reprints² also Dept. Botany³ also Institute of Applied Mathematics     

Limiting nutrient patchiness and its rôle in phytoplankton ecology

Limiting nutrient patchiness is examined as a factor affecting the community structure and species succession of natural phytoplankton communities held in ammonia limited continuous culture at a dilution rate of 0.3 day^?1. It was found that under a homogeneous distribution of the limiting nutrient members of genus Chaetoceros dominated and when ammonia was added daily (patchy distribution), Skeletonema dominated. Intermediate patchiness gave rise to an assemblage dominated by both Chaetoceros and Skeletonema. The nutrient uptake ability of each assemblage was determined three weeks after experiment initiation. Each assemblage was best able to optimize uptake of ammonia under its particular patchy nutrient regime. Optimization of a patchy environment took place by an increased maximal uptake rate (V_max) while optimization of a homogeneous environment appeared to take place by increased substrate affinity (i.e., low K_s).This study demonstrates that limiting nutrient patchiness can alter the relative abundance of populations within a community based on each population's ability to exploit the limiting resource under a particular degree of patchiness. We also show that coexistence of two populations might be expected due to the patchiness of a single limiting nutrient. The importance of patchiness in relation to other factors which determine community structure is discussed.     

Primary productivity,phytoplankton and limiting nutrient factors in labrador lakes

The primary productivity of some lakes and reservoirs in western Labrador was measured by the ¹⁴C method in order to determine the range of productivities and the effects of impoundment. No primary productivity data previously existed for this part of Canada. Both the primary productivity and standing crops of phytoplankton were found to be low in a newly impounded lake but later rose to levels greater than in surrounding natural lakes. In nutrient enrichment experiments, carbon was never found to be limiting but phosphorus stimulated primary productivity when added alone or in combination with nitrogen.     

Grazer control and nutrient limitation of phytoplankton biomass in two Australian reservoirs

1. Grazer and nutrient controls of phytoplankton biomass were tested on two reservoirs of different productivity to assess the potential for zooplankton grazing to affect chlorophyll/phosphorus regression models under Australian conditions. Experiments with zooplankton and nutrients manipulated in enclosures, laboratory feeding trials, and the analysis of in-lake plankton time series were performed. 2. Enclosures with water from the more productive Lake Hume (chlorophyll a = 3–17.5 μg l^–1), revealed significant zooplankton effects on chlorophyll a in 3/6, phosphorus limitation in 4/6 and nitrogen limitation in 1/6 of experiments conducted throughout the year. Enclosures with water from the less productive Lake Dartmouth (chlorophyll a = 0.8–3.5 μg l^–1), revealed significant zooplankton effects in 5/6, phosphorus limitation in 5/6 and nitrogen limitation in 2/6 of experiments. 3. While Lake Hume enclosure manipulations of the biomass of cladocerans (Daphnia and Diaphanosoma) and large copepods (Boeckella) had negative effects, small copepods (Mesocyclops and Calamoecia) could have positive effects on chlorophyll a. 4. In Lake Hume, total phytoplankton biovolume was negatively correlated with cladoceran biomass, positively with copepod biomass and was uncorrelated with total crustacean biomass. In Lake Dartmouth, total phytoplankton biovolume was negatively correlated with cladoceran biomass, copepod biomass and total crustacean biomass. 5. In both reservoirs, temporal variation in the biomass of Daphnia carinata alone could explain more than 50% of the observed variance in total phytoplankton biovolume. 6. During a period of low phytoplankton biovolume in Lake Hume in spring–summer 1993–94, a conservative estimate of cladoceran community grazing reached a maximum of 0.80 day^–1, suggesting that Cladocera made an important contribution to the development of the observed clear-water phase. 7. Enclosure experiments predicted significant grazing when the Cladocera/Phytoplankton biomass ratio was greater than 0.1; this threshold was consistently exceeded during clear water phase in Lake Hume. 8. Crustacean length had a significant effect on individual grazing rates in bottle experiments, with large Daphnia having highest rates. In both reservoirs, mean crustacean length was negatively correlated with phytoplankton biovolume. The observed upper limit of its variation was nearly twice as high compared to other world lakes.     

Sources of uncertainty in assessment of marine phytoplankton communities

Characterisation of phytoplankton communities is important for classification of the ecological status of marine waters. In order to design a monitoring programme, it is important to know what degree of variation in the measurements occur at each level (water body, station and sample), so that resources can be spent in a way that maximise the precision of the measured parameters. Seven European water bodies were sampled to assess the variation in pigment concentrations and population densities attributed to water body, station and sample levels. It was found that the main proportion of the variation between pigment measurements was explained by the variation between stations (12–91% of variation) followed by the variation between water bodies (0–89% of variation). For measurements of population density, the main proportion of the variation between densities of cells recorded was explained by the variation between the taxonomists counting the samples (61%), whilst the main proportion of the variation between numbers of taxa recorded was explained by the variation between water bodies (83%). When the cell density of the nine dominant classes were analysed separately, the main proportion of variation was explained at the water body level for all but two class.     

The effect of nutrient addition on the phytoplankton community of an oligotrophic lake

This investigation was performed during a 5-yr period (1974–1978) in the oligotrophic Lake Langvatn, Central Norway. In 1975 and 1976 the lake was enriched with a commercial fertilizer, In 1975 increase in phytoplankton biomass was first recorded more than three weeks after the fertilization started, despite a near fivefold increase in the primary production after fertilizer application. The mean seasonal biomass increased from c. 3500 mg wet weight m⁻² in 1974 to 4400 mg in 1975. In 1976 the biomass increased to near 9600 mg ⁻² and the seasonal primary production to 49.0 g C m⁻² (22.2 g C in 1975), despite a reduction in the nutrients added. Chrysophytes constituted the largest share of the seasonal algal biomass in all years, but in fertilization periods cryptophytes dominated in 1975 and diatoms, chlorophytes and cryptophytes in 1976. The highest biomass turnover rate was recorded during a period of cryptophyte dominance. The different biomass and production development in the fertilization years may be explained by a change in the consumer level.     

Response of three phytoplankton bioassay techniques in experimental ponds of known limiting nutrient

In a controlled enrichment study of eight experimental ponds, results from the batch bioassay, primary productivity incubation bioassay, and chemostat techniques for measuring limiting factors of phytoplankton algae were compared to the change in the natural system with nutrient addition. In the ponds, rapid and dramatic increase in both phytoplankton biomass and primary productivity upon the addition of nitrogen and phosphorus fertilizer offered conclusive evidence that these nutrients were limiting in the control ponds to which no nutrients were added. Both the batch bioassay and chemostat techniques clearly indicated nitrogen and possibly phosphorus as the limiting factors; however, the primary productivity incubation bioassay technique showed no increase in ¹⁴C uptake with addition of these nutrients. A species- and/or nutrient-specific time lag between nutrient uptake and increased carbon fixation is suggested to explain the failure of the technique to yield positive results within the 4-hour incubation period used.     

Description of phytoplankton and nutrient in spring in the western North Atlantic Ocean

Abundance of diatoms in the northern Sargasso Sea in springhas been observed in a 4-year period between 1981 and 1985 andthen again in 1987. A theory is presented that if stratificationblocks get-through of deep nutrients to the surface, as in thesouthern Sargasso, then in-situ cycling via non-diatoms andammonia dominates. The theory is that if in-situ cycling doesnot dominate, as northward, then stratification does not blockget-through of deep nutrients to the surface, which, in combinationwith the marked growth capacity of diatoms, produce an abundanceof cells. This abundance consumes nitrate, phosphate and silicateat the surface and ultimately this nutrient-depleted water arrivesin the southern Sargasso via the gyre circulation. Thus thenutrient-depleted water of the western Atlantic gyre is consideredto be primarily achieved only by diatoms, in spring and in thenorthern Sargasso. A transect during April 13–16, 1985,north from the Virgin Islands along 63°–64°W showsa great change in the diatom species plus Emiliania huxleyifrom negligible concentrations in the southern half of the sectionto large concentrations in the northern half. This unevennessof distribution can be rephrased by indicating that these speciesare narrow-niched with regard to nutrient availability. Theother species, coccolithophores and dinoflagellates, are evenlydistributed at low concentrations south-north (though some diminishslightly) and so they are broad-niched with respect to nutrient.High species diversity depends on broadness of nutrient nicheand equilibrium conditions of hydrographically stable water(i), implying that when non-equilibrium conditions occur (ii),when niche breadth is less (iii) or both (iv), then diversityis less marked, (i) is confirmed by broad-niched species southward,(ii) is confirmed by broad-niched species northward, (iii) isconfirmed by narrow-niched species southward and (iv) by narrow-nichedspecies northward.