DIEL PERIODICITY OF PHOTOSYNTHESIS AND CELL DIVISION COMPARED IN THALASSIOSIRA WEISSFLOGII (BACILLARIOPHYCEAE)1

Diel patterns of photosynthesis and cell division were examined in Thalassiosira weissflogii Grun. (clone Actin) grown in nitrogen-limited cyclostat culture. Ammonia (NH₄+) was either supplied continuously or as a daily pulse to cultures grown in constant light or in a light: dark cycle. When either nitrogen or light was supplied periodically, both cell division and photosynthetic capacity were periodic. When both nitrogen and light were supplied periodically, cell division was coupled to the N-pulse whereas periodicities of photosynthetic capacity were modified but remained coupled to the light-dark cycle. Diel oscillations in photosynthesis were i) largely independent of cellular pigmentation and ii) similar for light-limiting and saturating irradiances. Periodicity in photosynthetic capacity also persisted following transfer of non-dividing batch cultures to constant light. Results suggest that photosynthesis but not cell division was coupled to a circadian clock in T. weissflogii. A circadian rhythm of photosynthesis may optimize carbon assimilation in phytoplankton exposed to intermittent nutrient supply by ensuring that maximum photosynthetic capacity occurs during the day.     

The simultaneous assimilation of ammonium and l-arginine by the marine diatom Phaeodactylum tricornutum Bohlin

Nitrogen-replete cells of Phaeodactylum tricomutum Bohlin assimilated ammonium and the amino acid l-arginine simultaneously. Arginine was taken up at rates expected to supply at least 30% of the cells' requirement for nitrogen; arginme-carbon mainly entered protein but, when uptake was in darkness, ≈40% was respired. Cells grown in a 12:12 h light:dark cycle with ammonium as the sole nitrogen source took up ammonium throughout the growth cycle, whereas cells grown with the addition of arginine took up little ammonium during the dark phase. The uptake of ammonium over the course of the cycle was reduced by 30% when arginine was present. Cells grown with arginine as the sole nitrogen source took up the amino acid at the rate required for growth. In contrast, cells grown on ammonium, while growing at the same rate as those on arginine, assimilated nitrogen at twice the rate. Cells grown with both sources of nitrogen present, took up arginine at the same rate as before, but more of the arginine-carbon was respired (60% as compared with 40% when ammonium was absent). The uptake of ammonium was reduced by 30%, but the total nitrogen assimilation again exceeded immediate requirements. A high uptake rate of arginine was indicative of cells assimilating ammonium only; a low uptake rate of ammonium during the dark phase of growth was indicative of cells assimilating arginine. It is not known whether the findings with P. tricomutum are applicable to other marine phytoplankton. If they are, arginine may be of greater significance as a natural source of nitrogen for phytoplankton than is generally thought.     

In situ nutrient enrichment experiment in the Bohai and Yellow Sea

Nutrient concentrations and N : P ratios have changed significantlyin the past 40 years in the Bohai and Yellow Sea. How do thesechanges influence or contribute to the growth of phytoplankton?Nutrient enrichment experiments were conducted in 1998 and 1999to shed light on which was the first nutrient to limit algalgrowth and uptake rates of nitrogen and phosphorus. Significantvariance analysis, together with nutrient concentration andratio, demonstrated that phosphorus was the first nutrient tolimit the growth of phytoplankton in the Laizhou Bay (SouthBohai); nitrogen was the first, whilst phosphate might be thepotential, nutrient to limit the growth of phytoplankton inthe West Yellow Sea; the Central Yellow Sea was oligotrophicand any one of nitrogen, phosphorus and silicon would limitthe growth of phytoplankton; and silicon was confirmed not tolimit the growth of phytoplankton, although the silicate concentrationdecreased acutely, in the Laizhou Bay. Moreover, the ratio ofnitrogen to phosphorus in phytoplankton uptake was smaller thanthat in seawater, which suggested that phosphorus was preferentiallyused before nitrogen by the phytoplankton. The preference ofphosphorus over nitrogen indicates a further limitation of phosphorusin the Laizhou Bay, and increases the high possibility thatphosphorus, rather than nitrogen, is the first nutrient to limitthe growth of phytoplankton in the West Yellow Sea. Half-saturationconstants (K_s) of dissolved inorganic nitrogen (DIN) and phosphatein the Bohai and Yellow Sea were 1.80 µM and 0.13 µM,respectively. Compared with other sea areas, the K_s value ofDIN in the Bohai and Yellow Sea was located at the high endof the spectrum.     

Daily changes of uptake of inorganic carbon and nitrogen,and their relation to phytoplankton blooms in late spring-early summer in Lake Nakanuma,Japan

Daily changes of inorganic carbon and nitrogen uptake were measured in May in 1986 in Lake Nakanuma, Japan. Uptake of inorganic carbon and ammonium in the light-bottle experiments in the 1 m layers, showed daily changes similar to chlorophyll a changes, though the uptake activities peaked before chlorophyll a peaks (phytoplankton blooms) appeared. Potential growth rates of phytoplankton and observed growth rates were calculated from the uptake rates and chlorophyll a changes. The potential growth rates did not always correspond to the observed growth rates. The potential growth rates did not correlate with the loss rates. The correlation between the observed growth rates and the loss rates was better. These results suggest that though the increase of uptake activities may be necessary for occurrence of phytoplankton blooms, loss processes may affect the occurrence of blooms.     

Asynchronous vertical migration and bimodal distribution of motile phytoplankton

Some motile phytoplankton have the capability to exploit deepsources of nutrients in a vertical migration cycle: photosynthesisin the near-surface layer, transit to depth, uptake of the limitingnutrient and transit back to the surface layer. If all foursteps can be completed within 24 h, then migrations can be synchronizedto the day/night cycle to maximize photosynthetic efficiency.Alternatively, if physiological, behavioral or environmentalfactors make it impossible for the cycle to be completed in24 h, then migration may be asynchronous. Many observationsof phytoplankton reveal bimodal vertical distributions of organisms,with maxima near the surface and the nutricline. We demonstratehow bimodal vertical distributions of phytoplankton may be symptomaticof asynchronous vertical migration using a Lagrangian Ensemblenumerical model. We simulate vertical migration of the dinoflagellateAlexandrium fundyense in conditions similar to those in theGulf of Maine, where bimodal distributions of A. fundyense havebeen observed. Migration is regulated by internal nutritionalstate—organisms swim down toward the nitracline when depletedof nitrogen, and return to the surface after nutrient uptake.We test the sensitivity of the results to growth rate, nitrogenuptake rate and swimming speed, and find that organism distributionscan be bimodal or unimodal depending on conditions. Finally,we develop an analytical estimate for population distributionbased on organism characteristics and nutricline depth.     

Plankton and primary production in aquatic systems of the Central Amazon basin

• 1. Three aquatic environments of the Central Amazon Basin were sampled during the low water phase: Rio Solimo˜es (Amazon), Rio Negro and Lago Junauaca´.
• 2. In the nutrient-rich Rio Solimo˜es, primary productivity averaged 0.063 g C/m² per day, but net growth of phytoplankton is impossible because of the depth, poor light penetration and current-driven turbulence.
• 3. In the chemically-poor Rio Negro, a daily cycle of ΣCO₂ was caused by photosynthetic demand in surface waters, and primary productivity averaged 0.19 g C/m² per day.
• 4. In Lago Janauaca´. a lake bordering the Solimo˜es, nutrients were low in concentration and intensively recycled; large phytoplankton populations produced up to 3.5 g C/m² per day (average = 2.2).
• 5. The 10 m annual change in water level of the Amazon River provides and drains most of the water in lakes along the river. During storage of river water in the lakes the particulates settle and the dissolved inorganic nutrients are converted to organic forms by intense phytoplankton growth. These lakes may be a major source of organic matter for the river ecosystem.

     

Seasonal pattern in nutrient limitation and grazing control of the phytoplankton community in a non-stratified lake

1. The relative importance of zooplankton grazing and nutrient limitation in regulating the phytoplankton community in the non-stratified Lake Kvie, Denmark, were measured nine times during the growing season.
2. Natural phytoplankton assemblage bioassays showed increasing importance of nutrient limitation during summer. Growth rates at ambient nutrient concentrations were continually below 0.12 per day, while co-enrichment with nitrogen (N) and phosphorus (P) to above concentration-saturated conditions enhanced growth rates from May to the end of July.
3. Stoichiometric ratios of important elements in seston (C : N, C : P, N : P), in lake water (TN : TP), in external loading (TN : TP) and in internal loading (DIN : DIP) were measured to determine whether N or P could be the limiting nutrient. TN : TP molar ratio of both lake water, benthic fluxes and external loading suggested P limitation throughout the growing season. However, seston molar ratios suggested moderate P-deficiency only during mid-summer.
4. Abundance and community structure of the zooplankton varied considerably through the season and proved to be important in determining the responses of algal assemblages to grazing. High abundance of cladocerans and rotifers resulted in significant grazing impact, while cyclopoid copepods had no significant effect on the phytoplankton biomass.
5. Regeneration of ammonium and phosphate by zooplankton were periodically important for phytoplankton growth. A comparison of nutrient regeneration by zooplankton with nutrient inputs from sediment and external sources indicated that zooplankton may contribute significantly in supplying N and P for the growth of phytoplankton.     

Temporal and vertical dynamics of phytoplankton net growth in Castle Lake, California

The impact of nutrient additions, zooplankton grazing and light intensity on phytoplankton net growth with depth and season was studied with five microcosm experiments in meso-oligotrophic, subalpine Castle Lake, California, during the period of summer stratification in June-September 1994. The incubations (4 day) were performed at 5 m intervals from the surface to the bottom using natural phytoplankton and zooplankton assemblages, with enrichments of phosphorus and nitrogen. The phytoplankton community was only limited by nutrients in the upper 5 m (epilimnion), as indicated by change in chlorophyll concentration. Nutrient enrichments had the greatest effect on the phytoplankton net growth in June and July. High light inhibited the phytoplankton net growth at the surface. Low light intensities limited phytoplankton at 20 m and below, and at the end of the growing season already around 10-15 m. A deep chlorophyll maximum in the hypolimnion in June-August was not limited by either light or nutrients. The results showed variation in grazers' impact on phytoplankton. These results suggest the importance of nutrient limitation only in the epilimnion with light inhibition at the surface, light limitation in the hypolimnion, and varying impact of zooplankton grazing in influencing the development of the phytoplankton in Castle Lake.      

Diel variation of phytoplankton functional groups in a subtropical reservoir in southern Brazil during an autumnal stratification period

A knowledge of diel variation and the vertical distribution of phytoplankton communities may contribute to a better understanding of the driving factors of key species. Applying functional-group classification provides important information on the causes of species selection in the pelagic community. The diel variation of phytoplankton functional groups was analysed during an autumnal stratification period with the aim of understanding their changes in the vertical position related to light, mixing regime and grazing pressure. Phytoplankton and zooplankton communities were sampled every 4 h during a 24-h period in a vertical profile in a subtropical meso-eutrophic reservoir. Strong stratification during a 24-h cycle and a mixed clear epilimnion with partial atelomixis marked the autumn season in the Faxinal reservoir, southern Brazil. The highest phytoplankton densities and biomass were found during the second part of the day, a general pattern reported in the literature, and may be explained by zooplankton dynamics. During the 24-h cycle, phytoplankton functional groups lacking a self-regulating capacity and those able to regulate their vertical position were vertically segregated in the lake. The diel behaviour of both groups was driven by the mixing regime (including atelomixis), light and zooplankton grazing pressure.     

A PERSPECTIVE ON PHOTOSYNTHESIS IN THE OLIGOTROPHIC OCEANS: HYPOTHESES CONCERNING ALTERNATE ROUTES OF ELECTRON FLOW1

Many regions of the open, oligotrophic oceans are depleted of nutrients, especially nitrogen and iron. The biogenesis and the functioning of the photosynthetic apparatus may be specialized and tailored to the various marine habitats. In this minireview, we discuss some new findings with respect to photosynthetic processes in the oceans. We focus on findings that suggest that some cyanobacteria may route electrons derived from the splitting of H₂O to the reduction of O₂ and H⁺ in a water‐to‐water cycle, and that other cyanobacteria that fix nitrogen during the day are likely missing PSII and enzymes involved in the fixation of inorganic carbon. Both of these proposed “variant” forms of photosynthetic electron flow provide new insights into ways in which marine phytoplankton satisfy their energetic and nutritive requirements.     

Phytoplankton growth rates in the Ross Sea, Antarctica, determined by independent methods: temporal variations

The development of the seasonal phytoplankton bloom in the Ross Sea was studied during two cruises. The first, conducted in November-December 1994, investigated the initiation and rapid growth of the bloom, whereas the second (December 1995-January 1996) concentrated on the bloom's maximum biomass period and the subsequent decline in biomass. Central to the understanding of the controls of growth and the summer decline of the bloom is a quantitative assessment of the growth rate of phytoplankton. Growth rates were estimated over two time scales with different methods. The first estimated daily growth rates from isotropic incorporation under simulated in situ conditions, including ¹⁴C, ¹⁵N and ³²Si uptake measurements combined with estimates of standing stocks of particulate organic carbon, nitrogen and biogenic silica. The second method used daily to weekly changes in biomass at selected locations, with net growth rates being estimated from changes in standing stocks of phytoplankton. In addition, growth rates were estimated in large-volume experiments under optimal irradiances. Growth rates showed distinct temporal patterns. Early in the growing season, short-term estimates suggested that growth rates of in situ assemblages were less than maximum (relative to the temperature-limited maximum) and were likely reduced due to low irradiance regimes encountered under the ice. Growth rates increased thereafter and appeared to reach their maximum as biomass approached the seasonal peak, but decreased markedly in late December. Differences between the major taxonomic groups present were also noted, especially from the isotopic tracer experiments. The haplophyte Phaeocystic antarctica was dominant in 1994 throughout the growing season, and it exhibited the greatest growth rates (mean 0.41 day^-1) during spring. Diatom standing stocks were low early in the growing season, and growth rates averaged 0.100 day^-1. In summer diatoms were more abundant, but their growth rates remained much lower (mean of 0.08 day^-1) than the potential maximum. Understanding growth rate controls is essential to the development of predictive models of the carbon cycle and food webs in Antarctic waters.      

Ecological Significance of Nitrogen Cycling by Tubificid Communities in Shallow Eutrophic Lakes of the Danube Delta

Importance of tubificid populations on nitrogen cycle in two categories of shallow eutrophic lakes in the Danube Delta was quantitatively assessed for the 1992-1993 period. The structure of the primary producers in the studied lakes was used to discriminate between the two categories:(i) lakes dominated by macrophytes (A₁) and (ii) lakes dominated by phytoplankton (A₂). In both categories tubificid worms represented important fraction of the entire benthic community (35 and 32%, respectively, as number of individuals). They influence the sediment-water exchange of nutrients. The main processes involved are excretion of nutrients and their continuous release from sediments by molecular diffusion or through channels created by bioturbation. Inorganic nitrogen released from bottom sediments may regulate nitrogen load in the water body and thus, phytoplankton production. In 1992-1993, nitrogen stocks in tubificid biomass accounted for 5.3% in A₁ lakes and 15.6% in A₂ lakes of the amount stocked in phytoplankton, and only for 1.2 and 2.9% respectively, of the nitrogen load in water body. Nitrogen excretion rates ranged between 60.52 and 153.74 mg N m^–2 year^–1, and release rates from sediments between 378.26 and 960.87 mg N m^–2 year^–1, the lowest values being recorded for A₂ category. Differences are related to tubificid biomass, structure and abundance of primary producers and to nutrient load in different ecosystems. Ratios between release rate of inorganic nitrogen by tubificid worms and sedimentation rate of organic nitrogen in the two categories of lakes were 8.3 and 6.4% respectively. Contribution of nitrogen released daily from sediments to the dissolved inorganic nitrogen load in the water column was less than 0.5%. However, in A₁ and A₂ lakes, the released nitrogen had a potential to sustain 24.74 and 8.01%, respectively, of the annual phytoplankton production. These values suggest the significance of tubificids in keeping the eutrophication process at a high level, especially during the periods when nitrogen is the main limiting factor for phytoplankton production.     

DIFFERENCES IN GROWTH AND PHYSIOLOGY OF MARINE SYNECHOCOCCUS (CYANOBACTERIA) ON NITRATE VERSUS AMMONIUM ARE NOT DETERMINED SOLELY BY NITROGEN SOURCE REDOX STATE1

The preference of phytoplankton for ammonium over nitrate has traditionally been explained by the greater metabolic cost of reducing oxidized forms of nitrogen. This “metabolic cost hypothesis” implies that there should be a growth disadvantage on nitrate compared to ammonium or other forms of reduced nitrogen such as urea, especially when light limits growth, but in a variety of phytoplankton taxa, this predicted difference has not been observed. Our experiments with three strains of marine Synechococcus (WH7803, WH7805, and WH8112) did not reveal consistently faster growth (cell division) on ammonium or urea as compared to nitrate. Urease and glutamine synthetase (GS) activities varied with nitrogen source in a manner consistent with regulation by cellular nitrogen status via NtcA (rather than by external availability of nitrogen) in all three strains and indicated that each strain experienced some degree of nitrogen insufficiency during growth on nitrate. At light intensities that strongly limited growth, the composition (carbon, nitrogen, and pigment quotas) of WH7805 cells using nitrate was indistinguishable from that of cells using ammonium, but at saturating light intensities, cellular carbon, nitrogen, and pigment quotas were significantly lower in cells using nitrate than ammonium. These and similar results from other phytoplankton taxa suggest that a limitation in some step of nitrate uptake or assimilation, rather than the extra cost of reducing nitrate per se, may be the cause of differences in growth and physiology between cells using nitrate and ammonium.     

PHYTOPLANKTON LIPIDS: INTERSPECIFIC DIFFERENCES AND EFFECTS OF NITRATE,SILICATE AND LIGHT-DARK CYCLES1

The lipid content of various phytoplankton species was measured in response to nitrogen and silicon limitation and over the cell cycle in synchronized cultures. In a survey of 30 species it was found that during log-phase growth, green algae contained an average of 17.1% total lipids (% of total dry weight), whereas diatoms contained an average of 24.5%. Nitrogen deprivation for 4 to 9 days resulted in 2- to 3-fold increases in the lipid content of green algae, whereas both increases and decreases were noted in diatoms, depending on the species. The greatest lipid content measured in the study was 72% in Monallantus salina (strain GSB Sticho) which had been deprived of nitrogen for 9 days. Nitrate replenishment in a nitrogen starved culture of Oocystis polymorpha Groover & Bold showed that the excess cellular lipids do not rapidly disappear during recovery, until cell division occurs. A silicate deprivation experiment with Cyclotella cryptica Reimann, Lewin & Guillard (strain 7c) showed an increase in the total cellular lipid fraction from. 30 to 42% of dry weight within 6 h of the onset of silicon limitation, while the mass of lipid material per cell doubled within 12 h. The total lipid fraction in O. polymorpha was found to remain constant over the cell cycle in synchronized cultures regardless of the light regime. The data presented provided the first internally consistent study of phytoplankton lipids for a wide range of species and several growth conditions.     

Seasonal variations of size-fractionated phytoplankton along the salinity gradient in the York River estuary, Virginia (USA)

The dynamics of phytoplankton size structure were investigatedin the freshwater, transitional and estuarine zones of the YorkRiver over an annual cycle. The contribution of large cells(microplankton, >20 µm) to total concentrations ofchlorophyll a increased downstream during winter, whereas thatof small cells (nanoplankton, 3–20 µm; picoplankton,<3 µm) increased downstream during summer. In the freshwaterregion, the contribution of micro phytoplankton to total concentrationsof chlorophyll a was significant during warm seasons (springand summer) but not during colder seasons (winter), whereasthe contribution of small-sized cells (especially picoplankton)increased during cold seasons. Temperature, light and high flushingrate appear to control phytoplankton community structure inthe freshwater region. In the transitional region, nano-sizedcells dominated the phytoplankton population throughout allseasons except during the spring bloom (April) when the chlorophylla concentration of micro phytoplankton increased. Size structurein the transitional region is most likely regulated by lightavailability. In the mesohaline region, nano- and pico-sizedcells dominated the phytoplankton population during the summerbloom, whereas micro-sized cells dominated during the winterbloom. Factors controlling phytoplankton community size structurein the mesohaline zone may be riverine nitrogen input, temperatureand/or advective transport from up-river. Based on these results,the spatial and seasonal variations in size structure of phytoplanktonobserved on the estuarine scale may be determined both by thedifferent preferences for nutrients and by different light requirementsof micro-, nano- and picoplankton. The results suggest thatanalyses of phytoplankton size structure are necessary to betterunderstand controls on phytoplankton dynamics and to bettermanage water quality in river-dominated, estuarine systems.     

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.     

Phosphorus and nitrogen limitation of phytoplankton growth in eutrophic Lake Inba, Japan

Lake Inba is one of the most eutrophic lakes in Japan. In this study, field sampling and nutrient enrichment bioassays were conducted to determine the seasonal patterns of nutrient limitation for phytoplankton growth in this lake. Phytoplankton biomass increased significantly with the additions of phosphorus (P) on almost all sampling dates, indicating P limitation of phytoplankton growth from spring to autumn. However, nitrogen (N) limitation was also observed during summer (i.e., 19 August). On 10 August, a typhoon struck Lake Inba. After this event, dissolved inorganic nitrogen (DIN) and phosphorus concentrations increased, probably because of increased river discharge. At the same time, phytoplankton growth in the control treatment became relatively high, with the addition of neither P nor N stimulating the growth. However, 10 days after the typhoon, the phytoplankton growth rate in the control treatment decreased, with only the addition of N having a significant positive effect on phytoplankton growth. N limitation during summer is caused by the low concentrations of DIN, as well as changes in the N:P ratio due to allochthonous nutrient loads. These results indicate that a reduction of both P and N input is necessary to control phytoplankton blooms in Lake Inba.     

A mathematical model of the nutrient dynamics of phytoplankton in a nitrate-limited environment

Insofar as saturation kinetics are applicable to the growth of phytoplankton in laboratory experiments and to growth in nature, the computer modeling of intracellular nutrient partitioning in populations of cells can lead to better understanding of the dynamics of natural populations. A three-compartment mathematical model was developed to represent a phytoplankton population having the capability to store nitrogen in a nitrate-limited environment. Parameters were estimated by fitting the model to data from two chemostat experiments reported by Caperon (1968). The model was used to simulate growth dynamics observed in chemostat and batch experiments. The model demonstrated the changes which may occur in the nitrogenous constituents of a phytoplankton population with time and environmental conditions. The model also demonstrates three phenomena which have been observed in field and laboratory experiments but which are not represented by the customary Monod model: (1) uptake rates may significantly exceed not growth rates, (2) high growth rates may be encountered at very low environmental nitrate concentrations, and (3) the ratio of internal nitrogen to population size may change significantly during a study period. It is suggested that the amount of nitorgen in storage may be used as an indicator of the physiological state of a monospecific population. Parameters for the one-compartment Monod model were estimated by customary methods form data generated by the three-compartment model. It was shown that difficulties encountered in estimating the yield coefficient and the decay coefficient may be attributed to the intracellular storage phenomenon. It was also demonstrated that the one-compartment Monod model was inadequate to accurately represent population growth in chemostat experiments when intracellular storage is a significant factor.     

Nitrogen effects on the pelagic food web are modified by dissolved organic carbon

Global environmental change has altered the nitrogen (N) cycle and enhanced terrestrial dissolved organic carbon (DOC) loadings to northern boreal lakes. However, it is still unclear how enhanced N availability affects pelagic food web efficiency (FWE) and crustacean zooplankton growth in N limited boreal lakes. Here, we performed in situ mesocosm experiments in six unproductive boreal Swedish lakes, paired across a DOC gradient, with one lake in each pair fertilized with N (2011: reference year; 2012, 2013: impact years). We assessed how zooplankton growth and FWE were affected by changes in pelagic energy mobilization (PEM), food chain length (phytoplankton versus bacterial production based food chain, i.e. PP:BP), and food quality (seston stoichiometry) in response to N fertilization. Although PP, PEM and PP:BP increased in low and medium DOC lakes after N fertilization, consumer growth and FWE were reduced, especially at low DOC—potentially due to reduced phytoplankton food quality [increased C: phosphorus (P); N:P]. At high DOC, N fertilization caused modest increases in PP and PEM, with marginal changes in PP:BP and phytoplankton food quality, which, combined, led to a slight increase in zooplankton growth and FWE. Consequently, at low DOC (<12 mg L^?1), increased N availability lowers FWE due to mismatches in food quality demand and supply, whereas at high DOC this mismatch does not occur, and zooplankton production and FWE may increase. We conclude that the lake DOC level is critical for predicting the effects of enhanced inorganic N availability on pelagic productivity in boreal lakes.     

Nutrient pulsing as a regulator of phytoplankton abundance and community composition in Galveston Bay, Texas

Galveston Bay, Texas, is a large shallow estuary with a watershed that includes 60% of the major industrial facilities of Texas. However, the system exhibits low to moderate (2-20 μg l⁻¹) microalgal biomass with sporadic phytoplankton blooms. Both nitrogen (N) and phosphate (P) limitation of phytoplankton growth have been proposed for the estuary. However, shifts between N and P limitation of algae growth may occur due to annual fluctuations in nutrient concentrations. The primary goal of this work was to determine the primary limiting nutrient for phytoplankton in Galveston Bay. Nutrient addition bioassays were used to assess short-term (1-2 days) phytoplankton responses (both biomass and community composition) to potentially limiting nutrients. The experimental bioassays were conducted over an annual cycle using natural water collected from the center to lower part of the estuary. Total phytoplankton biomass increased in the nitrate (10 μM) additions in 11 of the 13 bioassays, but no significant increases were detected in the phosphate (3 μM)-only additions. Bioassay results suggest that the phytoplankton community was usually not phosphate limited. All major groups increased in biomass following nitrate additions but diatoms increased in biomass at a faster rate than other groups, shifting the community composition toward higher relative abundance of diatoms. The results of this study suggest that pulsed N input events preferentially favor increases in diatom biomass in this estuary. The broader implications of this study are that N pulsing events, primarily due to river discharge, play an important role in structuring the phytoplankton community in the Galveston Bay estuary.