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.     

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     

Relations between planktivorous fish abundance,zooplankton and phytoplankton in three lakes of differing productivity

Hydrobiologia - Water chemistry, phytoplankton, zooplankton and fish populations werestudied over several years in three shallow, non-stratified lakes withdiffering nutrient loadings and fish...     

Thermal stratification, nutrient dynamics, and phytoplankton productivity during the onset of spring phytoplankton growth in Lake Baikal, Russia

Lake Baikal, Russian Siberia, was sampled in July 1990 during the period of spring mixing and initiation of thermal stratification. Vertical profiles of temperature, dissolved nutrients (nitrate and soluble reactive phosphorus), phytoplankton biomass, and primary productivity were determined in an eleven-station transect encompassing the entire 636 km length of the lake. Pronounced horizontal variability in hydrodynamic conditions was observed, with the southern region of the lake being strongly thermally stratified while the middle and north basins were largely isothermal through July. The extent of depletion of surface water nutrients, and the magnitude of phytoplankton biomass and productivity, were found to be strongly correlated with the degree of thermal stratification. Horizontal differences likely reflected the contribution of two important factors: variation in the timing of ice-out in different parts of the lake (driving large-scale patterns of thermal stratification and other limnological properties) and localized effects of river inflows that may contribute to the preliminary stabilization of the water column in the face of intense turbulent spring mixing (driving meso-scale patterns). Examination of the relationships between surface water inorganic N and P depletion suggested that during the spring and early summer, phytoplankton growth in unstratified portions of the lake was largely unconstrained by nutrient supplies. As summer progressed, the importance of co-limitation by both N and P became more apparent. Uptake and regeneration rates, measured directly using the stable isotope ¹⁵N, revealed that phytoplankton in stratified portions of the lake relied primarily on NH₄ as their N source. Rates of NH₄ regeneration were in approximate equilibrium with uptake; both processes were dominated by organisms <2 µm. This pattern is similar to that observed for oligotrophic marine systems. Our study underscores the importance of hydrodynamic conditions in influencing patterns of biological productivity and nutrient dynamics that occur in Lake Baikal during its brief growing season.     

Primary production and phytoplankton in two small,polyhumic forest lakes in southern Finland

Primary production and phytoplankton in polyhumic lakes showed a very distinct seasonal succession. A vigorous spring maximum produced by Chlamydomonas green algae at the beginning of the growing season and two summer maxima composed mainly of Mallomonas caudata Iwanoff were typical. The annual primary production was ca. 6 g org. C · m^–2 in both lakes. The mean epilimnetic biomass was 1.1 in the first lake and 2.2 g · m^–2 (ww) in the second one. The maximum phytoplankton biomass, 14 g · m^–2, was observed during the vernal peak in May.     

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.     

Responses of epilimnetic phytoplankton to experimental nutrient enrichment in three small seepage lakes

This paper describes the responses of three epilimnetic phytoplanktoncommunities to experimental nitrogen and phosphorus enrichmentas compared to the phytoplankton community in a fourth, unmanipulated,lake. Increased nutrient inputs increased total phytoplanktonbiomass, primary productivity, chlorophytes, cryptomonads andspecies turnover rates in all three enriched lakes; cyanobacteriaincreased in two of the three enriched lakes. However, nutrientaddition also led to declines in previously dominant dinoflagellatesand chrysophytes, and in species diversity. At the species level,there were large changes in community composition from yearto year in both enriched and reference lakes, suggesting thatphytoplankton community composition is highly dynamic even inthe absence of enrichment. Overall, changes in total biomass,productivity and species diversity were consistent among theenriched lakes, while changes in species composition differeddue to variation in the physical, chemical and biotic environmentof each lake. This suggests that aggregated variates are moreuseful for quantitative prediction of nutrient effects, whilespecies responses can be used to signal qualitative differencesin environmental conditions among lakes. ³Present address: Department of Biological Sciences, DartmouthCollege, 6044 Gilman Laboratory, Hanover, NH 03755-3576, USA     

An experimental investigation of phytoplankton nutrient limitation in two contrasting low arctic lakes

We investigated whether phytoplankton communities in two lakes in SW Greenland were phosphorus or nitrogen limited. The study lakes have contrasting water chemistry (mean conductivities differ ten fold) and are located near Kangerlussuaq, SW Greenland (~67°N, 51°W). A microcosm nutrient enrichment experiment was performed in June 2003 to determine whether nitrate or phosphate addition stimulated phytoplankton growth. Samples were analysed for species composition, biomass, and alkaline phosphatase activity (APA). Initially, both lakes had extremely low total phosphorus but high total nitrogen concentrations and high APA, suggesting that the phytoplankton were phosphorus limited prior to the start of the experiment. The phytoplankton composition and biomass (mainly Ochromonas spp.) responded to phosphate but not to nitrate addition. In both lakes, chlorophyll a increased significantly when phosphate was added. Furthermore, APA was significantly lower in the two lakes when phosphate was added compared to the control and the nitrogen addition treatment. The dominance of mixotrophic phytoplankton and high DOC values suggest that these lakes may be regulated by microbial loop processes.     

30 Seasonal control of phytoplankton biomass and productivity in coastal british columbia lakes and reservoirs

The algal class Chlorarachniophyceae is comprised of a small group of unicellular eukaryotic algae that are often characterized by an unusual amoeboid morphology. This morphology is hypothesized to be the result of a secondary endosymbiosis in which a green alga was engulfed as prey by a nonphotosynthetic amoeba or amoebaflagellate. Whereas much is known about the phylogenetic relationships of individual chlorarachniophytes to one another, and to possible ancestral host organisms in the genera Cercomonas and Heteromita , little is known about their physiology, particularly that of their lipids. In an initial effort to characterize the lipids of this algal class, seven organisms were examined for their fatty acid and sterol composition. These included Bigelowiella natans, Chlorarachnion globusum, Chlorarachnion reptans, Gymnochlora stellata, Lotharella amoeboformis, Lotharella globosa , and Lotharella sp . Fatty acids associated with chloroplast-associated glycolipids, cytoplasmic membrane-associated phospholipids, and storage triglycerides were characterized. Glycolipid fatty acids were found to be of limited composition, containing principally eicosapentaenoic acid [20:5(n-3)] and hexadecanoic acid (16:0), which ranged in relative percentage from 67–90% and 10–29%, respectively, in these seven organisms. Triglyceride-associated fatty acids were found to be similar. Phospholipid fatty acid composition was more variable. The principal phospholipid fatty acids, 16:0 (25–32%) and a compound tentatively identified as docosapentaenoic acid [22:5(n-3)] (26–35%), were found along with a number of C18 and C20 fatty acids. All organisms contained two sterols as free sterols. These were tentatively identified as 24-ethylcholesta-5,22E-dien-3b-ol (stigmasterol; 70–95%) and 24-methylcholesta-5,22E-dien-3b-ol (brassicasterol; 5–30%).     

The upper limit of photosynthetic productivity by phytoplankton: evidence from Ethiopian soda lakes

Photosynthetic production by phytoplankton was studied in two Ethiopian soda lakes, alkalinity 51-67 m-equiv./l, with abundant blue-green algae. The deeper lake, L. Aranguadi, contained extremely dense crops composed almost entirely of SpiruUna (Oscillatoria, Arthrospira) platensis. Measurements of the spectral attenuation of light showed that the most penetrating component was displaced to the red spectral region, and estimates of the chlorophyll a content in umt area of the euphotic zone were often similar to the highest values (～200-300 mg/m²) expected on theoretical grounds. The vertical distribution of photosynthetic activity per unit water volume was of a typical pattern for phytoplankton, with light-inhibition often present, although the euphotic zones did not exceed 0.6 m in depth. From these profiles, computed rates of gross photosynthesis per unit area of lake surface reached a magnitude of 1.4–2.4 g O₂/m².h in both lakes. They were obviously severely limited by self-shading behaviour i n the algal populations. An appreciable inverse relationship between photosynthetic capacity and population density was only found in the densest populations of L. Aranguadi, with chlorophyll a content > 2000 mg/m³, where depressed rates might result from experimental artefacts in closed bottles. In this lake, two independent estimates of gross production, based on the analysis of diurnal changes in the open water, were as high as 43 and 57 g O₂/m². day. Diurnal changes of stratification in the two lakes are described and related to the controlling temperature (density) stratification. They include occasionally complete nocturnal deoxygenation in the deeper lake, evidence of heavy respiratory uptake. The high photosynthetic productivity is interpreted as dependent upon the coupling of high algal contents in the euphotic zone (^S) with high values of photosynthetic capacity (^max)- It is probably favoured by the tropical situation affecting temperature and illumination, by a surplus of dissolved inorganic phosphate, and especially by the considerable reserves of carbon dioxide in these soda lakes.     

Diel periodicity in phytoplankton productivity

Daily variation in phytoplankton productivity influences the dynamics and linkages between several large scale processes in aquatic ecosystems. As part of an opening address to the 5th International workshop for the Group for Aquatic Productivity (GAP), the daily patterns of variability in photosynthesis for different algal classes was introduced and accompanied by a discussion of the sources of environmental and endogenous regulation of repeating biological oscillations that occur in phytoplankton on timescales of one day. It is suggested that one way to develop a database that serves to sort and predict phytoplankton variability over the day may be to encourage the creation of a temporal library. Such a library would be comprised of temporally fixed maps of circadian clock-controlled rhythms for individual species, as well as temporally variable maps of diel periodicities that only can be defined for a selected set of environmental conditions.     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.     

Primary production and extracellular release by phytoplankton in some lakes of the Masurian Lake District, Poland.

The primary production and extracellular release by phytoplankton were measured at different depths of Miko?ajskie Lake, Ryńskie Lake and Be?dany Lake (Masurian Lake District, Poland). The release of dissolved organic compounds was found to be related to the rates of primary production. The percentage of extracellular release (PER) of organic matter in the photic zone showed an inverse relationship with particulate production and chlorophyll alpha concentration. The highest PER was determined in the epilimnion of the studied lakes, where also increased release was observed. The higher release of algal products of photosynthesis in the photic zone than in the profundal is probably caused by the inhibition of physiological activity of bacteria by antibacterial substances produced by algae. The substances released by algae in the profundal are taken up by aquatic bacteria which explains the lower release and PER measured.     

Primary production of phytoplankton and periphyton in two humic lakes of a South American wetland

Seasonal primary productivities of periphyton and phytoplankton were compared in Grande Lake (GL) and a relict oxbow lake (ROL) in winter 2006 and summer 2007. GL was free of floating plants on the sampling dates and covered over 80 and 100% of the ROL surface in winter and summer, respectively. The ¹⁴C assimilation technique was used to obtain the P–E curves of phytoplankton and periphyton on artificial substrata. The periphytic maximum photosynthetic rate (P _max) was higher in the ROL in winter and summer, being better adapted to low irradiances than those in the GL. Phytoplankton and periphytic algae were light-limited in the ROL in summer due to complete coverage by floating macrophytes. In summer, P _max and α values for periphyton in the ROL were higher than those for phytoplankton, and were even higher than in GL. In turn, P _max and α values for phytoplankton in Grande Lake were higher than those for periphyton due to improved light conditions and the presence of algae that were adapted to movement through the water column. These results suggest that the complete coverage by floating macrophytes restricted phytoplankton productivity and allowed the development of a periphytic community that was better adapted to low-light conditions.     

Spatial, seasonal and long-term variability of phytoplankton photosynthesis in lakes

A cross-system, worldwide approach has been used to ascertainthe spatial, seasonal and long-term variability of areal phytoplanktonphotosynthesis (PP) in lakes using published data sets. Also,the average fraction of annual PP occurring under ice is calculated.The lakes considered embrace a range of properties (depth, mixing,flushing rate, latitude and trophic status). The overall yearlyPP distribution is skewed to the left, suggesting the dominanceof low PP rates in the data set. When comparing lake types,no differences in average PP have been found among them. Inparticular, there are no clear areal PP differences among lakesof different trophic status on yearly, averaged basis, suggestingthat environmental limitations to PP also exist in lakes ofhigher trophic status. Volumetric-based PP can be better usedto outline PP-based trophic differences, but some degree ofoverlap is also apparent. Across all lake types (except in tropicallakes), the PP seasonal course experiences only one peak inthe year, but its timing is clearly different for each laketype. The seasonal variability of PP is lower in tropical lakes,as previously reported, but the variability of the other laketypes is roughly the same. Therefore, the effects of depth,mixing regime, flushing rate and nutrient status on PP seasonalityare difficult to ascertain since they appear to be counterbalancedby other more pervasive, local effects. Particularly, thereis no increase in temporal variability with the trophic statusof lakes, suggesting that PP seasonal control by physical variablesoverrides that of nutrients. Also, no significant relationshipbetween average PP and latitude has been found. Seasonal variabilityincreases as the yearly PP increases. On a relative basis, thereis a spatial gradient of seasonal variability of PP, which isweaker when seasonal variability of PP is considered in interyearcomparisons. Long-term (i.e. interannual) variability of PPis clearly related to increasing yearly averaged PP. Specifically,in temperate, stratifying lakes the seasonal time course ofPP is clearly different from that of phytoplankton biomass,suggesting an uncoupling of both variables as a result of differingP_max and losses throughout the year. On an average basis, environmentalvariables are poor predictors of areal daily PP, thereby implyingthat the interplay of factors is complex and changing throughoutthe year. PP under ice averages 10% of yearly PP, but its variabilityis high enough to make its measurement advisable.     

Detritivory and the stoichiometry of nutrient cycling by a dominant fish species in lakes of varying productivity

Little is known about the stoichiometry of nutrient cycling by detritivores. Therefore, we explored stoichiometric relationships in an omnivorous/detritivorous fish (gizzard shad, Dorosoma cepedianum) in three lakes that differed in productivity. Gizzard shad can feed on plankton and sediment detritus, but in all three lakes adult gizzard shad derived >98% of carbon (C) and phosphorus (P), and >90% of nitrogen (N) from sediment detritus, and the remainder from zooplankton.
Gizzard shad selectively consumed detritus with higher C, N and P concentrations than ambient lake sediments. Selective detritivory (i.e. the nutrient content of consumed detritus divided by the nutrient content of ambient detritus) was most pronounced in the lake with the lowest detrital nutrient concentrations. N and P cycling rates per fish were also consistently higher in this lake, in agreement with the prediction of stoichiometry theory that excretion rates should increase with food nutrient content. Among-lake differences in nutrient cycling rates were unrelated to inter-lake variation in fish body nutrient contents, which was minimal. The N:P ratio excreted was near Redfield (∼14:1) in all three lakes.
Stoichiometric analyses showed that the C:N and C:P ratios of sediment detritus were much higher (∼2.8×) than ratios of gizzard shad bodies, revealing substantial N and P imbalances between consumers and their food source. Gizzard shad alleviate N imbalance by selectively feeding on high N detritus (low C:N, high N:P), and apparently alleviate P imbalance by excreting nutrients at a higher N:P than that of their food or their bodies. Thus, this detritivore apparently regulates nutrient acquisition and allocation via both pre-absorption processes (selective feeding) and post-absorptive processes (differential N and P excretion).     

Effects of resource pulses on nutrient availability,ecosystem productivity,and temporal variability following a stochastic disturbance in eutrophic glacial lakes

Tropical macrophytes sold in the live garden trade are perceived as unlikely to invade temperate regions owing to climate mismatches. Here we study two tropical macrophytes (Pistia stratiotes and Eichhornia crassipes) not previously considered an invasion risk but which were recently discovered in the Great Lakes, and determine mechanisms that may be responsible for their continued presence including human introduction, reproduction through viable seeds and tolerance of winter conditions. Surveys conducted in 2011 and 2012 revealed recurrent presence of one or both species at some sites. Macrophytes in in situ enclosures failed to survive winter conditions, with plant health declining progressively prior to mortality. Water hyacinth seeds were field-collected, identified using Sanger sequencing, and germinated at 28°C with or without scarification. Germination was highest for scarified versus non-scarified seeds. Human introduction was observed at two sites, one involving both species, the other only water hyacinth. These species likely persist through a combination of annual reintroduction (both species) and possibly by production of viable seed (water hyacinth). Macrophytes, particularly water hyacinth, that were not previously viewed as a threat to the Great Lakes owing to environmental incompatibility may need to be reassessed.     

Effects of glaciers on nutrient concentrations and phytoplankton in lakes within the Northern Cascades Mountains (USA)

We compared nitrate concentrations, phytoplankton biomass, and phytoplankton community structure in lakes fed by glacier melt and snowmelt (GSF lakes) and by snowmelt only (SF lakes) within North Cascades National Park (NOCA) in Washington State, USA. In the U.S. Rocky Mountains, glacier melting has greatly increased nitrate concentrations in GSF lakes (52–236 µg NO₃–N L^?1) relative to SF lakes (1–14 µg NO₃–N L^?1) and thereby stimulated phytoplankton changes in GSF lakes. Considering NOCA contains approximately one-third of the glaciers in the continental U.S., and many mountain lakes that receive glacier meltwater inputs, we hypothesized that NOCA GSF lakes would have greater nitrate concentrations, greater phytoplankton biomass, and greater abundance of nitrogen-sensitive diatom species than NOCA SF lakes. However, at NOCA nitrate concentrations were much lower and differences between lake types were small compared to the Rockies. At NOCA, nitrate concentrations averaged 13 and 5 µg NO₃–N L^?1 in GSF and SF lakes, respectively, and a nitrate difference was not detectable in several individual years. There also was no difference in phytoplankton biomass or abundance of nitrogen-sensitive diatoms between lake types at NOCA. In contrast to the Rockies, there also was not a significant positive relationship between watershed percent glacier area and lake nitrate at NOCA. Results demonstrate that biogeochemical responses to global change in Western U.S. mountain lake watersheds may vary regionally. Regional differences may be affected by differing nitrogen deposition, climate, geology, or microbial processes within glacier environments, and merit further investigation.     

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.