Growth rate of alpine phytoplankton assemblages from contrasting watersheds and N‐deposition regimes exposed to nitrogen and phosphorus enrichments

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Transplant experiments. Growth-limiting factors for diatoms and bluegreen algae in Norwegian lakes

Population densities and total phosphorus concentrations in samples from different lakes of south-eastern Norway were determined. In addition some transplant experiments with dilute phytoplankton populations were carried out. A laboratory batch culture method was used.The diatoms studied may be divided into three ecological groups based on their cell densities and total phosphorus concentrations in the samples. This classification was supported by the experimental results. Cyclotella spp., Asterionella formosa and Tabellaria fenestrata did not grow or had low growth rates above pH 9. Synedra cf. acus and Fragilaria crontonensis had often high growth rates within the pH 9–10 range, but were not able to grow at pH values above 10. High pH-values had no effect on the growth rate of Oscillatoria. Oscillatoria, Synedra and Stephanodiscus were severely growth-limited in filtered water from oligotrophic lakes. Maximum growth rates of all the populations studied were often obtained after addition of phosphate and chelated iron (FeEDTA) in combination to filtered water samples from oligotrophic/mesotrophic lakes.     

Determination of available phosphorus for phytoplankton populations in lakes and rivers of southeastern Norway

A biotest method with diluted phytoplankton populations was used to determine external concentrations of available phosphorus in water samples with high concentrations of inorganic seston from River Rømua. RP (total molybdate reactive P measured on unfiltered samples) was approximately the P fraction available for Synedra cf. acus, Asterionella formosa and Oscillatoria agardhii. In filtered samples RP_F was the available concentration of P.The ratio RP:RP_F may give valuable information on the ratio between total available P (including available P in seston) and available P in filtered water. Different filter types may give different RP: RP_F ratios. The ratio RP:RP_F was often high during spring and autumn in River Rømua and the lakes studied. During the period June–September RP:RP_F 1 in most of the lakes and periodically in River Rømua.     

Nutrient limitation of bacterioplankton and phytoplankton in humic lakes in northern Sweden

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L^–1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), respectively. A third lake was unfertilized and served as a control. In addition to this lake fertilization experiment, data from different regional surveys were used to assess the role of different limiting factors.
2. The P fertilization had no effects on bacterioplankton or phytoplankton, while phytoplankton were significantly stimulated by N fertilization. Inorganic nutrient limitation of bacterioplankton was a function of DOC concentration in water of the investigated region and nutrient-limited bacteria were found only in lakes with DOC concentrations less than around 15 mg L^–1
3. The fertilization experiments demonstrated that the DOC-rich experimental lakes contained a bioavailable pool of P that was not utilized to its full potential under natural conditions. The overall mobilization of energy (bacterioplankton plus phytoplankton) in the experimental lakes was restricted by lack of inorganic N.     

Inter-annual,seasonal and spatial variability in nutrient limitation of phytoplankton production in a river impoundment

We characterize seasonal and spatial patterns in phytoplankton abundance, production and nutrient limitation in a mesotrophic river impoundment located in the southeastern United States to assess variation arising from inter-annual differences in watershed inputs. Short-term (48 h) in situ nutrient addition experiments were conducted between May and October at three sites located along the longitudinal axis of the lake. Nutrient limitation was detected in 12 of the 18 experiments conducted over 2 years. Phytoplankton responded to additions of phosphorus alone although highest chlorophyll concentrations were observed in enclosures receiving combined (P and N) additions. Growth responses were greatest at downstream sites and in late summer suggesting that those populations experience more severe nutrient limitation. Interannual variation in nutrient limitation and primary production corresponded to differences in the timing of hydrologic inputs. Above average rainfall and discharge in late-summer (July–October) of 1996 coincided with higher in-lake nutrient concentrations, increased production, and minimal nutrient limitation. During the same period in 1995, discharge was lower, nutrient concentrations were lower, and nutrient limitation of phytoplankton production was more pronounced. Our results suggest that nutrient limitation is common in this river impoundment but that modest inter-annual variability in the timing of hydrologic inputs can substantially influence seasonal and spatial patterns.     

Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.     

Seasonality in Southern Hemisphere freshwater phytoplankton assemblages

Seasonal climatic cycles induce corresponding fluctuations in phytoplankton abundance and productivity at all latitudes, the magnitude of these fluctuations tending to increase with distance from the Equator. In equatorial regions seasonality is dependent on prevailing wind and rainfall patterns while annual temperature fluctuations exert increasing control over seasonal events at higher latitudes. The small annual temperature range of equatorial aquatic systems increases their sensitivity to localized climatic events which can bring about diel changes that exceed normal month-to-month variations. Long-term hydrological cycles with a periodicity greater than one year can also cause dramatic changes in equatorial and tropical aquatic systems leading to greater unpredictability.The factors regulating seasonal patterns of phytoplankton abundance and species composition in equatorial and low-latitude temperate regions of the Southern Hemisphere are examined and compared with similar features in the Northern Hemisphere. Despite the striking diversity of phytoplankton populations and the wide variety of habitats they occupy, seasonal succession follows a common sequence controlled, successively, by physical, chemical and biotic factors. This permits a high degree of predictability in the environmental conditions promoting growth of different taxa.Examination of Southern Hemisphere data indicates that, at class level, phytoplankton successional sequences in Southern Hemisphere aquatic systems are in agreement with the successional paradigm formulated for northern tropical and temperate latitudes. Diatoms characterize early successional episodes and these are followed by chlorophytes, and finally blue-green algae. Extreme habitat modification (e.g. hypertrophy, salinity) tends to lead to dominance of the habitat by a single taxon, often represented by a single species. Predictions of within-taxon species succession in phytoplankton assemblages are far less precise.     

Nutrient limitation of phytoplankton and periphyton growth in upland lakes

SUMMARY 1. Thirty small upland lakes in Cumbria, Wales, Scotland and Northern Ireland were visited three times between April and August 2000. On each occasion water chemistry was measured and phytoplankton bioassays were performed in the laboratory to assess growth‐rate and yield limitation by phosphorus and nitrogen. In addition, yield limitation of periphyton growth was investigated twice, in situ, using nutrient‐diffusing substrata. 2. Over the whole season the percentage frequency of P, N and co‐limitation was 24, 13 and 63%, respectively, for phytoplankton rate limitation and 20, 22 and 58%, respectively, for phytoplankton yield limitation. 3. A clear response of periphyton yield to nutrient additions was found in 75% of all cases and of these, co‐limitation was most common (54%). Average percentage frequency for P and N limitation was 26 and 20%, respectively. 4. Phytoplankton and periphyton showed seasonal changes in nutrient limitation within sites. In particular, co‐limitation became progressively more common as the season progressed. 5. The response of phytoplankton growth rate to ammonium and nitrate addition was identical, but ammonium was a slightly better source of nitrogen than nitrate for phytoplankton yield on 7% and for periphyton yield on 10% of the occasions. However, the magnitude of the effect was small. 6. The concentration of dissolved inorganic nitrogen (DIN) and the molar ratio of DIN to total dissolved phosphorus (TDP), appeared to be the main environmental factors controlling the extent of nitrogen or phosphorus limitation at a given site. Nitrogen limitation was more likely than phosphorus limitation where the DIN was <6.5 mmol m^?3 and the ratio of DIN : TDP was <53. Co‐limitation was the most likely outcome at a DIN concentration <13 mmol m^?3 and at a DIN : TDP molar ratio <250. Above these values phosphorus limitation was most likely. 7. The relatively high frequency of nitrogen limitation and co‐limitation at higher N : P ratios than previously reported, may result from the inability of nitrogen‐fixing cyanobacteria to thrive in these upland lakes where pH and the concentration of phosphorus tended to be low and where flushing rates tended to be high.     

The influence of mesozooplankton on phytoplankton nutrient limitation: a mesocosm study with northeast Atlantic plankton

We used marine phytoplankton from mesocosms seeded with different zooplankton densities to study the impact of mesozooplankton on phytoplankton nutrient limitation. After 7 d of grazing (copepod mesocosms) or 9 d (appendicularian mesocosms) phytoplankton nutrient limitation was studied by enrichment bioassays. After removal of mesozooplankton, bioassay bottles received either no nutrients, phosphorus or nitrogen alone, or a combination of nitrogen and phosphorus and were incubated for 2 d. Phytoplankton reproductive rates in the bottles without nutrient addition were calculated after correction for grazing by ciliates and indicated increasing nitrogen limitation with increasing copepod abundance. No nutrient limitation was found in the appendicularian mesocosms. The increase of nutrient limitation with increasing copepod density seems to be mainly the result of a trophic cascade effect: Copepods released nanoplankton from ciliate grazing pressure, and thereby enhanced nitrogen exhaustion by nanophytoplankton and reduced nitrogen excretion by ciliates. Nitrogen sequestration in copepod biomass, the mechanism predicted by the ecological stoichiometry theory, seems to have been a weaker effect because there was only little copepod growth during the experiment.     

Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

Interactive effect of warming,nitrogen and phosphorus limitation on phytoplankton cell size

Cell size is one of the ecologically most important traits of phytoplankton. The cell size variation is frequently related to temperature and nutrient limitation. In order to disentangle the role of both factors, an experiment was conducted to determine the possible interactions of these factors. Baltic Sea water containing the natural plankton community was used. We performed a factorial combined experiment of temperature, type of nutrient limitation (N vs. P), and strength of nutrient limitation. The type of nutrient limitation was manipulated by altering the N:P ratio of the medium (balanced, N and P limitation) and strength by the dilution rate (0% and 50%) of the semicontinuous cultures. The negative effect of temperature on cell size was strongest under N limitation, intermediate under P limitation, and weakest when N and P were supplied at balanced ratios. However, temperature also influenced the intensity of nutrient imitation, because at higher temperature there was a tendency for dissolved nutrient concentrations to be lower, while the C:N or C:P ratio being higher…higher at identical dilution rates and medium composition. Analyzing the response of cell size to C:N ratios (as index of N limitation) and C:P ratios (as index of P limitation) indicated a clear dominance of the nutrient effect over the direct temperature effect, although the temperature effect was also significant.     

Nutrient limitation of phytoplankton in a central Arizona reservoir

The influence of added nutrients nitrogen, phosphorus and carbon on the phytoplankton of a small recreational reservoir in central Arizona was investigated during the summer, 1974. Polyethylene bags were used to isolate lake water and the natural populations for the addition of nitrogen, phosphorus and carbon individually and in combination. A large increase in phytoplankton numbers, extractable chlorophyll, pH and dissolved oxygen occurred only in bags to which both nitrogen and phosphorus were added, suggesting that both nitrogen and phosphorus levels were limiting to the primary producers. Little alteration in species composition resulted from the addition of the above nutrients.     

太平洋中西部海域浮游植物营养盐的潜在限制

2009年8月至9月期间在太平洋西部N1站和中部N2站进行现场营养盐加富培养实验。结果显示:N1站,浮游植物生物量对N或者P添加都有较强的响应,其中N+P+Si组和N+P组浮游植物长势迅速,叶绿素a从初始的0.03μg/L分别达到2.12μg/L和1.83μg/L,同时P先于N和Si之前被耗尽;说明N1站为N、P共同限制,P是首要限制因子。而N2站,浮游植物生物量仅对N、P共同添加有明显响应,N先于P和Si被浮游植物消耗殆尽。利用培养过程中营养盐比值变化推断,N1站浮游植物以低于Redfield ratio(16N∶1P)吸收N和P;而N2站浮游植物以高于Redfield ratio(16N∶1P)吸收N和P。这可能解释了太平洋西部的寡营养盐海域为潜在P限制,而在太平洋中部海域则为潜在N限制。     

The structure of phytoplankton assemblages in Sicilian reservoirs: some hyphoteses on the driving factors

Abstract

From the analysis of the results of several researches carried out into Sicilian reservoirs to investigate their phytoplankton assemblages, it was highlighted that there is not any clear relationship between the trophic spectrum and the structure and composition of phytoplankton. Only a general increase of total biomass is detectable as the trophic state rises up. Anyway, such increase may cause several secondary modifications in the physical and chemical environment that may act as multidimensional segregating factors for phytoplankton organisms. In these secondary modifications, coupled with the peculiar morphology and hydrology of the water bodies, have to be sought the mechanisms governing the structure of phytoplankton assemblages.     

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.     

Temporal variation in plankton assemblages and physicochemistry of Devils Lake, North Dakota

Seasonal and annual variation in biomass and structure of algal assemblages of hyposaline Devils Lake were examined in relation to turbidity, ambient concentrations of major ions, trace elements and nutrients, and the standing crop of herbivores. Lake level declined during the early years of study, but rose markedly in subsequent years as historically large volumes of water flowed into this hydrologically-closed basin. Winter algal assemblages were dominated (in biomass) most years by small, non-motile chlorophytes ( Choricystis minor, Kirchneriella lunaris or Dunaliella sp.), or Euglena sp. in the most saline sub-basin. Spring assemblages were dominated by diatoms (Stephanodiscus cf. minutulus, Surirella peisonis, Cyclotella meneghiniana and Entomoneis paludosa were especially prominent) or chlorophytes ( C. minor) until the lake level rose. C. minor abundances then declined in spring assemblages and diatoms ( Stephanodiscus cf. agassizensis and S. niagarae; E. paludosa in the more saline sub-basins) dominated. The potential for nitrogen-deficient conditions for phytoplankton growth was evidenced most summers and early autumns by consistently high concentrations of reactive-P relative to inorganic-N and blooms of the N-fixing cyanophyte Aphanizomenon flos-aquae; Microcystis aeruginosa typically was a co-dominant (>30% of biomass) in these assemblages. Pulses of diatoms ( S. cf. agassizensis and C. meneghiniana) occurred in summers following unusually prolonged periods of calm weather or large water inflows. Physical (irradiance, turbulence) and chemical (major nutrients) variables were the primary factors associated with phytoplankton growth. Transparency and major nutrient concentrations accounted for more of the annual variation in phytoplankton structure than did salinity. Seasonal abundance patterns of the dominant zooplankton (the copepod Diaptomus sicilis; the cladocerans Ceriodaphnia quadrangula, Chydorus sphaericus, Daphnia pulex and Diaphanosoma birgei; and the rotifers Brachionus spp., Filinia longiseta, Keratella cochlearis and K. quadrata) also indicated variation in algal populations related to grazing. This revised version was published online in July 2006 with corrections to the Cover Date.     

Micronutrient and phosphorus limitation of phytoplankton abundance in Gem Lake,Sierra Nevada,California

Nutrient addition experiments conducted during the ice-free seasons of 1983 and 1984 in Gem Lake, an alpine lake in the Sierra Nevada mountains of California, indicate that algal biomass is limited by phosphorus, in combination with iron or copper. Phosphorus additions were always required to stimulate growth, but did not do so when phosphorus was the only nutrient added. Simultaneous additions of phosphorus and iron resulted in increased levels of chlorophyll, particulate carbon, particulate nitrogen and particulate phosphorus. Simultaneous additions of phosphorus and copper resulted in increases in chlorophyll, particulate nitrogen and particulate phosphorus, not in particulate carbon. Neither iron nor copper by itself stimulated growth.Particulate N : P ratios from all seasons in Gem Lake suggest that simultaneous micronutrient and phosphorus limitation exists throughout the summer, when nutrient and biomass levels remain low; limitation by phosphorus alone may appear in the fall and spring, when biomass and major ion concentrations increase dramatically.     

Growth rates of phytoplankton under fluctuating light

• 1 The effect of light fluctuations on the growth rates of four species of freshwater phytoplankton was investigated. Experimental light regimes included constant irradiance and fluctuations of a step function form, with equal proportion of high (maximum of 240 µmol photons m^‐2 s^‐1) and low light (minimum of 5 µmol photons m^‐2 s^‐1) (or dark) in a period. Fluctuations of 1, 8 and 24‐h periods were imposed over several average irradiances (25, 50, 100 and 120 µmol photons m^‐2 s^‐1).
• 2 Growth rate responses to fluctuations were species‐specific and depended on both the average irradiance and the period of fluctuations. Fluctuations at low average irradiances slightly increased growth rate of the diatom Nitzschia sp. and depressed growth of the cyanobacterium Phormidium luridum and the green alga Sphaerocystis schroeteri compared to a constant irradiance.
• 3 Fluctuations at higher average irradiance did not have a significant effect on the growth rates of Nitzschia sp. and Sphaerocystis schroeteri (fluctuations around saturating irradiances) and slightly increased the growth rates of the cyanobacteria Anabaena flos‐aquae and Phormidium luridum (when irradiance fluctuated between limiting and inhibiting levels).
• 4 In general, the effect of fluctuations tended to be greater when irradiance fluctuated between limiting and saturating or inhibiting levels of a species growth‐irradiance curve compared to fluctuations within a single region of the curve.
• 5 The growth rates of species under fluctuating light could not always be predicted from their growth‐irradiance curves obtained under constant irradiance. When fluctuations occur between limiting and saturating or inhibiting irradiances for the alga and when the period of fluctuations is long (greater than 8 h), steady‐state growth‐irradiance curves may be insufficient to predict growth rates adequately. Consequently, additional data on physiological acclimation, such as changes in photosynthetic parameters, may be required for predictions under non‐constant light supply in comparison to constant conditions.

     

Uses of alkaline phosphatase activity in evaluating phytoplankton community phosphorus deficiency

The phosphorus (P) deficiency status of phytoplankton communities was measured using the physiological indicator, alkaline phosphatase activity (APA) and nutrient-addition growth bioassays in field sampled from four northeastern Minnesota lakes and the far western arm of Lake Superior. Phosphorus additions generally reduced APA, while other treatments increased activity. Samples receiving nitrogen (N) and P increased APA after a long lag period. P-addition bioassays of Lake Superior were consistent with phytoplankton P limitation and variations in APA indicated potential seasonal and spatial changes in P deficiency status. The results suggest that APA reliably reflected the phytoplankton P status, but may not provide sufficient information when N or NP limitation is present.