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.     

Primary production and phytoplankton composition in relation to DOC input and bacterioplankton production in humic Lake Örträsket

1. The biomass and production of picophytoplankton, large phytoplankton and heterotrophic bacterioplankton were measured in humic Lake Örträsket, northern Sweden during four consecutive summers.
2. High flow episodes, carrying fresh dissolved organic carbon (DOC) into the lake, always stimulated heterotrophic bacterial production at the expense of primary production. Primary production never exceeded bacterial production for approximately 20 days after such an episode had replenished epilimnial DOC. We suggest that allochthonous DOC is an energy source that stimulates bacterioplankton that, because of their efficient uptake of inorganic nutrients, are then able to outcompete phytoplankton. After the exhaustion of readily available DOC, phytoplankton were able to dominate epilimnion production in Lake Örträsket.
3. Biomass production was higher when dominated by phytoplankton than by bacterioplankton, despite a similar utilization of nutrients in the epilimnion throughout the summer. We propose that different C : N : P ratios of bacterioplankton and phytoplankton permit the latter to produce more carbon (C) biomass per unit of available inorganic nutrients than bacterioplankton.     

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.     

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.     

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.     

Relationships between picophytoplankton and environmental variables in lakes along a gradient of water colour and nutrient content

SUMMARY 1. Biomass and production of picophytoplankton, phytoplankton and heterotrophic bacterioplankton were measured in seven lakes, exhibiting a broad range in water colour because of humic substances. The aim of the study was to identify environmental variables explaining the absolute and relative importance of picophytoplankton. In addition, two dystrophic lakes were fertilised with inorganic phosphorus and nitrogen, to test eventual nutrient limitation of picophytoplankton in these systems.
2. Picophytoplankton biomass and production were highest in lakes with low concentrations of dissolved organic carbon (DOC), and DOC proved the factor explaining most variation in picophytoplankton biomass and production. The relationship between picophytoplankton and lake trophy was negative, most likely because much P was bound in humic complexes. Picophytoplankton biomass decreased after the additions of P and N.
3. Compared with heterotrophic bacterioplankton, picophytoplankton were most successful at the clearwater end of the lake water colour gradient. Phytoplankton dominated over heterotrophic bacteria in the clearwater systems possibly because heterotrophic bacteria in such lakes are dependent on organic carbon produced by phytoplankton.
4. Compared with other phytoplankton, picophytoplankton did best at intermediate DOC concentrations; flagellates dominated in the humic lakes and large autotrophic phytoplankton in the clearwater lakes.
5. Picophytoplankton were not better competitors than large phytoplankton in situations when heterotrophic bacteria had access to a non-algal carbon source. Neither did their small size lead to picophytoplankton dominance over large phytoplankton in the clearwater lakes. Possible reasons include the ability of larger phytoplankton to float or swim to reduce sedimentation losses and to acquire nutrients by phagotrophy.     

Biotests with phytoplankton assemblages. Growth limitation along temporal and spatial gradients

The development of eight different species (populations) along temporal and vertical gradients in several lakes was studied. Many populations had an exponential growth phase and a decline phase. The growth rate was often high during the exponential phase. Some species, e.g. Oscillatoria spp. and Synedra cf. acus, often also had a long stationary phase. The growth rate and the sinking rate of these populations were often very low. Laboratory batch experiments with dilute phytoplankton populations were carried out to estimate the degree of growth limitation (L) for different populations sample from different lakes during the three growth phases. L was always low and often zero for populations initially in the exponential phase and always high for populations initially in the decline phase. The biotests also gave results that can help to explain the vertical distribution of Oscillatoria or Asterionella in three lakes investigated. The results indicate that the growth rates and the development of the populations were dependent on the external chemical and physical conditions. The transition between the different growth phases seemed often to be dependent on the external nutrient conditions. P, N, Si and Fe were probably the most growth-limiting nutrients. The growth rate of some diatoms was probably limited directly or indirectly at high pH. Laboratory biotests with natural populations may give valuable information on the growth-properties of different populations in the lakes. The biotests should, however, be carried out in combination with chemical and physical measurements and quantitative determinations of population densities.     

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.     

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.     

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.     

Clay turbidity and the relative production of bacterioplankton and phytoplankton

A comparison of phytoplankton with bacterioplanktonproduction as each ismodified by high concentrations of suspended clays ispresented. High clayturbidity caused light-limition of water columnphytoplankton production.However, the clay combined with DOC to form aggregateswhich supportedbacterioplankton production. Consequently,bacterioplankton production wasrelatively high at 42% of total small particleproduction in this lake.Bacterioplankton abundance and biomass was stronglycorrelated withphytoplankton chlorophyll a for most of the lake. Because of the association ofbacterioplankton with the clay-organic aggregates, DOCwas not a good predictorof bacterioplankton abundance or production. POC(primarily OC associatedwith clay) was correlated with bacterioplanktonabundance over most of thelake. Bacteria production was substrate limited asshown by much greaterbiomass-specific production at smaller bacteriapopulation sizes. Multipleregression analysis showed that specificbacterioplankton production wasprimarily governed by POC and secondarily by rates ofphytoplanktonproduction. Thus clay, because of light limitedphytoplankton production,negatively impacts bacterioplankton production yet atthe same time facilitatiesbacterial production by concentrating OC with theformation of the clay-organicaggregate.     

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。     

Atmospheric nitrogen deposition has caused nitrogen enrichment and eutrophication of lakes in the northern hemisphere

We compiled chemical data and phytoplankton biomass (PB) data (chlorophyll a ) from unproductive lakes in 42 different regions in Europe and North America, and compared these data to inorganic nitrogen (N) deposition over these regions. We demonstrate that increased deposition of inorganic N over large areas of Europe and North America has caused elevated concentrations of inorganic N in lakes. In addition, the unproductive lakes in high N deposition areas had clearly higher PB relative to the total phosphorus (P) concentrations illustrating that the elevated inorganic N concentrations has resulted in eutrophication and increased biomass of phytoplankton. The eutrophication caused by inorganic N deposition indicates that PB yield in a majority of lakes in the northern hemisphere is (was) limited by N in their natural state. We, therefore, suggest that P limitation largely concerns lakes where the balance between N and P has been changed because of increased anthropogenic input of N.     

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.     

Ambiguous climate impacts on competition between submerged macrophytes and phytoplankton in shallow lakes

1. Shallow lakes may switch from a state dominated by submerged macrophytes to a phytoplankton‐dominated state when a critical nutrient concentration is exceeded. We explore how climate change may affect this critical nutrient concentration by linking a graphical model to data from 83 lakes along a large climate gradient in South America. 2. The data indicate that in warmer climates, submerged macrophytes may tolerate more underwater shade than in cooler lakes. By contrast, the relationship between phytoplankton biomass [approximated by chlorophyll‐a (chl‐a) or biovolume] and nutrient concentrations did not change consistently along the climate gradient. In warmer climates, the correlation between phytoplankton biomass and nutrient concentrations was overall weak, especially at low total phosphorus (TP) concentrations where the chl‐a/ TP ratio could be either low or high. 3. Although the enhanced shade tolerance of submerged plants in warmer lakes might promote the stability of their dominance, the potentially high phytoplankton biomass at low nutrient concentrations suggests an overall low predictability of climate effects. 4. We found that near‐bottom oxygen concentrations are lower in warm lakes than in cooler lakes, implying that anoxic P release from eutrophic sediment in warm lakes likely causes higher TP concentrations in the water column. Subsequently, this may lead to a higher phytoplankton biomass in warmer lakes than in cooler lakes with similar external nutrient loadings. 5. Our results indicate that climate effects on the competitive balance between submerged macrophytes and phytoplankton are not straightforward.     

The light : nutrient ratio in lakes: a test of hypothesized trends in bacterial nutrient limitation

The light-to-nutrient hypothesis explores how the balance between energy (as light energy) and nutrients (as total phosphorus) shapes aquatic ecosystem structure and process. The balance of energy and nutrients is thought to regulate ecosystem structure and process such that, in a "high" light-to-nutrient environment, bacteria would probably be driven towards phosphorus (P) limitation, whereas, in a "low" light-to-nutrient environment, bacteria would be driven towards carbon (C) limitation. We assessed the growth limitation of bacteria in two reservoirs of the southern U.S.A. using a mortality-corrected dilution-growth approach. We compared the frequency of P and C growth limitation with the intralake variation in the light-to-nutrient environment. As a metric of the light-to-nutrient environment, we used the ratio of the mean light in the surface mixed layer ( I _m) to the total phosphorus concentration ( TP ). In each lake, bacterial growth was more often P-limited when the I _m :  TP ratio was above the median ratio than below. We believe our data provide the first evidence supporting this aspect of the light-to-nutrient hypothesis.     

Light climate and phytoplankton photosynthesis in maritime Antarctic lakes

The responses of phytoplankton populations to seasonal changes in radiation flux in two Antarctic lakes with extensive winter ice-cover are described. A phytoplankton capable of photosynthesis was found throughout the year in both systems. During winter, low incident radiation combined with thick layers of snow and ice prevented in situ photosynthesis becoming detectable. The beginning of spring was marked by a reduction in snow cover which resulted in a considerable increase in surface penetrating radiation. Planktonic algae rapidly adapted to utilise these increased levels efficiently, though they still showed characteristics of strong shade adaptation.Loss of ice cover at the start of the short open water period further increased the radiation levels and a summer population developed which was much less shade adapted. Saturation and photoinhibition effects were widespread during this period as the algae proved unable to utilise high radiation levels efficiently. They were however effective at the radiation fluxes prevalent in the lower part of the rapidly circulating water columns.     

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.