In situ measurements confirm the seasonal dominance of benthic algae over phytoplankton in nearshore primary production of a large lake

1. Despite the recognition of its importance, benthic primary production is seldom reported, especially for large lakes. We measured in situ benthic net primary production by monitoring flux in dissolved inorganic carbon (DIC) concentration in benthic incubation chambers, based on continuous measurements of CO_2(aq) flux, alkalinity, and the temperature‐dependent dissociation constants of carbonic acid (K₁ and K₂). This methodology has the advantages of monitoring net primary production directly as change in carbon, maintaining continuous water recirculation, and having sufficient precision to detect change in DIC over short (i.e. 15 min) incubations, even in alkaline waters. 2. Benthic primary production on Cladophora‐dominated rocky substrata in western Lake Ontario was measured biweekly. Maximum biomass‐specific net photosynthetic rates were highest in the spring (2.39 mgC g Dry Mass^?1 h^?1), decreased to negative rates by early summer (?0.76 mgC g DM^?1 h^?1), and exhibited a regrowth in late summer (1.98 mgC g DM^?1 h^?1). 3. A Cladophora growth model (CGM), previously validated to predict Cladophora biomass accrual in Lake Ontario, successfully simulated the seasonality and magnitude of biomass‐specific primary production during the first cohort of Cladophora growth. Averaged over this growing season (May–Aug), mean areal net benthic production at the estimated depth of peak biomass (2 m) was 405 mg C m^?2 d^?1. 4. We measured planktonic primary production in proximity to the benthic study and constructed a depth‐resolved model of planktonic production. Using the CGM, benthic primary production was compared with planktonic primary production for the period May–Aug. Net benthic production from the shoreline to the 12 m contour (1–2 km offshore) equalled planktonic production. Closer to shore, benthic primary production exceeded planktonic primary production. Failure to account for benthic primary production, at least during abundant Cladophora growth, will lead to large underestimates in carbon and nutrient flows in the nearshore zone of this Great Lake.     

The maximum quantum yield of phylopiankton photosynthesis in situ

Light-limited photosynthetic carbon incorportion is expectedto be directly proportional to the scalar quantum irradiance.The proportionality constant is , where _mis the maximum quantum yield (mol C Einstein^–1 absorbed)and $$\stackrel{\¯}{{\hbox{ k }}_{\hbox{ c }}}$$ isthe mean spectral absorption coefficient (m² mg^–1 chla). Recent efforts to evaluate of in situphytoplankton photosynthesis are variously flawed. Lack of evidenceof proportionality and lack of correction of cosine to scalarirradiance are common deficiencies. Most data, as we interpretthem, indicate values in the range 0.0003 – 0.0006 mol C m² Einstein1 abs mg1 chl a. New determinationsin lrondequoit Bay, New York, lie in this range. Most estimatesof at depth have been about 0.010 m² mg^–1chl a. Similar values are being obtained for total particulatesfrom lrondequoit Bay; whether detritus contributes significantlyis not yet known. Published data, in our view, all point tovalues of m in situ in the range 0.03–0.07 mol C Einstein^–1abs. Published values >0.10 are almost certainly due to imprecisionor systematic error. ^*This paper is the result of a study made at the Group for AquaticPrimary Productivity (GAP) First International Workshop heldat the Limnological Institute, University of Konstanz, in April1982.     

Long‐term phytoplankton community changes in a deep subalpine lake: responses to nutrient availability and climatic fluctuations

1. In natural lakes, modifications in the species composition and abundance of phytoplankton communities may ultimately be responses to changes in nutrient availability and climatic fluctuations. Phytoplankton and associated environmental factors were collected at monthly intervals from the beginning of the 1990s to 2007 in the large subalpine Lake Garda (z_max = 350 m, V = 49 × 10⁹ m³). In this study period, the lake showed a slight and continuous increase of total phosphorus (TP) in the water column, up to concentrations of 18–20 μg P L^?1. This increase represented the last stage of a long‐term process of enrichment documented since the 1970s, when concentrations of TP were below or around 10 μg P L^?1. 2. At the community level, annual phytoplankton cycles underwent a unidirectional and slow shift mainly due to changes in the species more affected by the nutrient enrichment of the lake. After a first and long period of dominance by conjugatophytes (Mougeotia) and diatoms (Fragilaria), phytoplankton biomass in recent years was sustained by cyanobacteria (Planktothrix). Other important modifications in the development of phytoplankton were superimposed on this pattern due to the effects of annual climate fluctuations principally mediated by the deep mixing events at spring overturn and, secondarily, by temperature and thermal stability of the water column during the growing season. 3. Interannual variations in the stability and temperature of the water column appeared to influence the development of a few subdominant flagellates (dinophytes and cryptophytes). Nevertheless, the major impact of climate on phytoplankton was indirect, and mediated through the effects of winter climatic conditions on deep mixing dynamics. Winter climatic fluctuations proved to be a key element in a linked chain of causal factors including cooling of hypolimnetic waters, deep vertical mixing and epilimnetic nutrient replenishment. The process of fertilisation was measurable both for TP and dissolved inorganic nitrogen, although only the first had a large effect, reinforcing the seasonal growth of a few dominant groups. The degree of nutrient replenishment further increased the spring development of large diatoms and the increase of Planktothrix in summer and autumn. 4. Currently, changes in nutrient concentrations have the greatest effect on the phytoplankton community, while direct effects due to the interannual variations in the thermal regime are of secondary importance compared with the indirect effects mediated through deep water mixing and spring fertilisation. Overall, the results demonstrate that the consequences of climatic fluctuations and climate warming on phytoplankton communities need to be studied at different levels of complexity and integration, from the direct effects of temperature and thermal regime, to the indirect effects mediated by the physiographic characteristics of water bodies.     

In situ determination of sulfide turnover rates in a meromictic alpine lake

A push-pull method, previously used in groundwater analyses, was successfully adapted for measuring sulfide turnover rates in situ at different depths in the meromictic Lake Cadagno. In the layer of phototrophic bacteria at about 12 m in depth net sulfide consumption was observed during the day, indicating active bacterial photosynthesis. During the night the sulfide turnover rates were positive, indicating a net sulfide production from the reduction of more-oxidized sulfur compounds. Because of lack of light, no photosynthesis takes place in the monimolimnion; thus, only sulfide formation is observed both during the day and the night. Sulfide turnover rates in the oxic mixolimnion were always positive as sulfide is spontaneously oxidized by oxygen and as the rates of sulfide oxidation depend on the oxygen concentrations present. Sulfide oxidation by chemolithotrophic bacteria may occur at the oxicline, but this cannot be distinguished from spontaneous chemical oxidation.     

Response of lake phytoplankton communities to in situ manipulations of light intensity and colour

Phytoplankton preferences for light intensity and colour weredetermined in field experiments using coloured plexiglass cubessuspended at different depths in Heney Lake, Québec.Diatoms and green algae favoured intensities greater than 1%I_o (surface irradiance) contrary to dinoflagellates and otherflagellates that preferred lower intensity. Red radiation usuallyincreased the relative proportion of blue-greens, diatoms andgreen algae, whereas it reduced that of dinoflageilates. Wepropose that differential utilization of the light gradientallows certain phytoplankton taxa to partition the water column,thereby reducing potential competition. This is supported bythe general agreement between our findings and the known depthdistribution of algae in lakes.     

The effect of temperature on photosynthetic and respiratory electron transport system activity in the shallow and deep-living phytoplankton of a subalpine lake

SUMMARY. 1. The influence of temperature on in vivo photosynthetic and in vitro respiratory electron transport system (ETS) activity was determined over the season for the 3 m (warm-water) and a 20m (cold-water) phytoplankton communities in Castle Lake. The optimum temperature of photosynthesis at 3 m (X?=20.8°C) was significantly higher than the average optimum at 20 m (X?=14.8°C). 2. Seasonally, the photosynthetic temperature optimum increased when the blue-green alga Chroococcus limneticus Lemm. was present. The temperature characteristics of this organism were maintained even after it had settled into the cold water of the hypolimnion. 3. Temperature optima were not significantly different in experiments conducted under limiting or saturating photosynthetic photon flux densities (PPFD). 4. Short-term (1 h) preincubations with dissolved inorganic nitrogen (DIN) (?80 μg NH₄NO₃-N l^?1) had little effect on the temperature characteristics of photosynthesis while the longer (>24 h) incubations provided by a whole-lake epilimnetic DIN addition (?75 μg NH₄NO₃- N l^?1) significantly lowered the photosynthetic temperature optimum to 12.5°C. Once this epilimnetic DIN was depleted the optimum roseto25°C, a value higher than that present before the enrichment, which coincided with the growth of C limneticus. 5. Respiratory ETS activity usually began to inactivate between 19 and 20°C. However, when C. limneticus was abundant the inactivation temperature was often greater ihan 25°C. 6. The average energy of activation (E) and Q₁₀ value for the 3 m community (15.9 kcal mol^?1 and 2.6 respectively) were significantly higher than those at 20 m (14.2 kcal mol^?1 and 2.4 respectively). Seasonally, the highest E and Q₁₀ values of ETS activity occurred during the late-summer bloom of C. limneticus. 7. These results demonstrate that the epilimnetic and hypolimnetic phytoplankton communities in Castle Lake are physiologically distinct with regards to their temperature characteristics.     

Physical and chemical effects of macrograzers and micrograzers on enclosed,in situ phytoplankton in a Newfoundland lake

Grazing effects were evaluated by comparisons between phytoplankton populations in the lake water and in bicompartmental chambers, deployedin situ in a Newfoundland lake. Micrograzing effects were evaluated separately from both the physical effects of macrograzers and the chemical effects of macrograzer metabolites on phytoplankton assemblages in bicompartmental chambers. Macrograzer metabolites were able to pass through fine gauze between the compartments. Thus, the chemical effects of the macrograzers were observed in one compartment while the physical effects were the dominant treatment in the other. Micrograzer effects were investigated in chambers without added macrograzers.Densities of some of the desmid species were depressed by the physical effects of macrograzers. This is the first ecological study to demonstrate such a grazing effect on desmids. Some individual taxa, such asArthrodesmus triangularis var.rotundatus (Raciborski)comb. nov., andMesotaenium sp. Nägeli (Zygnemaphyceae), were augmented under the chemical effects of macrograzer metabolites. Evidence suggests that the desmids were not micrograzed. Grazing effects on desmids have not previously been studied in waters in which they were abundant.The Chlorophyceae, which were small species in Hogan's pond, and the microflagellates generally maintained their overall population densities in spite of the presence of macrograzers, however, abundances of some individual species within the assemblages changed in the presence of macrograzers. Micrograzers may also affect the species composition and abundance of the individual chlorophycean taxa.With the exceptions ofSynedra sp. Ehrenberg andTabellaria fenestrata var.lacustris Meister the diatoms were unaffected by the grazers. The Chrysophyceae were little affected by micrograzing and physical effects of macrograzers, in contrast to chemical effects of macrograzer metabolites that were evident for individual species.Several of the Cyanobacteria (Cyanophyceae) were augmented in the presence of micrograzers. Evidence indicates that they were not micrograzed. Some individual species, e.g.,Microcystis aeruginosa Kützing, were augmented by both physical and chemical effects of macrograzers.     

In situ sampling of interstitial water from lake sediments

A sampler with a relatively high resolution has been developed, which allows interstitial water to be obtained from lake sediments at well defined depths, without serious disturbance of sediment structure. Oxidation effects are excluded. Sampling time is in the order of a day. Installation requires little additional equipment. The instrument has been developed for use in shallow lakes.     

Dynamic status of primary production in lake yunoko,a small eutrophic subalpine lake in central Japan

Investigation was made of the primary productivity of Lake Yunoko. The concentrations of PO₄-P and NH₃-N were highest in summer. The total amount of chlorophylla in the entire water column varied from 45 mg/m² to 405 mg/m² during the year with the maximum values occurring in late spring and in December. These values are very high compared with those reported for other lakes in the same district. The standing crop of phytoplankton as organic carbon was approximately 50∼70% of the total particulate organic carbon in the lake with values as high as 80∼90% being obtained in the spring and autumn. The rate of phytosynthesis under saturated light conditions was 3.7 mg C/mg chla/hr in the case of phytoplankton of the sun type and 1.2∼1.6 mg C/mg chla/hr in those of the shade type. The daily gross primary production in the lake varied from a low of 134 mg C/m² to a high of 1,003 mg C/m² during the year, the mean value being 372 mg C/m². The total amount of daily microbial respiration varied from 177 mg C/m² in winter to 1,476 mg C/m² in summer. Thus, the daily balance of production and consumption of organic matter in the lake itself was concluded to be negative. The main factor contributing to the rectification of this imbalance of material budget seems to be the leaf litter coming from the forest surrounding the lake. In conclusion, Lake Yunoko is a very productive and at the same time, fairly heterotrophic lake. Special Project Research supported by the Ministry of Education on Dynamic State of Biosphere. This study was carried out as part of JIBP-PF.     

Interactions between phytoplankton and zooplankton in a fertilized lake

The limnology of an oligotrophic lake, Langvatn, situated in Trøndelag county in Central Norway, has been studied for five years (1974–1978). In two years, 1975 and 1976, the lake was fertilized with a general fertilizer to change feeding conditions for the zooplankton. Mean phytoplankton biomass in the epilimnion and primary production for the years (1974–1978) were 417, 618, 1370, 607 and 779 mg m⁻³ and 10.6, 22.2, 49.0, 26.8 and 17.7 g C m⁻² yr⁻¹, respectively. Cladocerans were the dominant herbivore group in 1974 and 1975 and Rotifera in the next three years. The main difference in the interaction between phytoplankton and zooplankton occurred when cladoceran dominance gave way to rotifer dominance. Heavy phytoplankton grazing by cladocerans in 1974 and 1975 stabilized the biomass and maintained it at a low level, which also resulted in a relatively low primary production. The rotifer-dominated community during the years 1976–1978 did not possess the ability to maintain a stable level of algal biomass. Primary production was also relatively high during these years.     

Cell death in lake phytoplankton communities

1. The fraction of living and dead phytoplankton cells in seven Florida lakes was assessed by using the cell digestion assay, a non‐staining membrane permeability test. The cell digestion assay is an effective method to analyse cell viability in complex natural phytoplankton communities. 2. The lakes examined ranged widely in phytoplankton abundance and community composition. The variability in the percentage of living cells (% LC) was high among the taxonomic groups forming the different phytoplankton communities, ranging from 19.7% to 98% LC. 3. All cells within single cyanobacteria filaments were determined to be either dead or alive, suggesting physiological integration of the cells within colonies. 4. Within each lake, the dominant taxa generally exhibited the highest proportion of living cells. A high proportion of living cells was found to be a characteristic of the different taxa forming the communities of eutrophic lakes. The average value for the % LC for all groups comprising the phytoplankton communities in each of the lakes ranged from 29.9 ± 7.2 to 80.4 ± 4.0 (mean ± SE) and varied strongly and positively with chlorophyll a concentration. 5. These results suggest phytoplankton cell death to be an important process structuring phytoplankton communities in lakes, particularly in oligotrophic ones.     

Effects of a massive swarm of ants on ammonium concentrations in a subalpine lake

On two occasions in August 1979, massive aerial swarms of Lasius alienus (Foerster) (Hymenoptera: Formicidae) occurred over Castle Lake, California. Each event resulted in the delivery of millions of ants to the lake surface. Consumption and subsequent excretion by fish, in addition to direct release from ants, produced a brief, but marked increase in water column ammonium levels. Castle Lake phytoplankton are often nitrogen limited and possess a high affinity for ammonium. The events and effects are described and discussed with respect to nutrient limitation in an unproductive ecosystem.     

In situ plankton community respiration measurements show low respiratory quotients in a eutrophic lake

Planktonic community respiration is an important carbon cycling process, typically quantified by converting measured values of dissolved O₂ consumption rates into CO₂ production rates assuming a respiratory quotient of 1 (RQ = CO₂ per O₂ by moles). However, the true variability in planktonic RQs between different aquatic ecosystems is poorly understood. We conducted in situ RQ measurements in a eutrophic lake dominated by algal-derived substances and found that RQs were significantly below 1. In fact, many RQ values were extremely low (0.2–0.6), below theoretical RQs for oxidation of algal organic matter substrates (0.7–0.8), suggesting that other factors than substrate control need to be considered to understand the RQ. This view was further supported by lack of correlations between RQ and microbial variables known to be strongly substrate dependent, including bacterial growth efficiency and the functional capacity of the bacterioplankton community to degrade different compounds. Based on the measured dynamics in methane and nutrient pools, we discuss that methane oxidation and nitrification likely occurred in the lake, contributing to the unusually low RQs. Our findings demonstrate that planktonic RQs in productive lakes can systematically be below 1, suggesting that CO₂ emissions from these lakes may currently be overestimated.     

A comparison of nitrogen and carbon metabolism in the shallow and deep-water phytoplankton populations of a subalpine lake: response to photosynthetic photon flux density

Rates of , and CO₂ assimilationby the organisms in the shallow and deepchlorophyll layers ofCastle Lake were measured over a gradient of photosyntheticphoton flux densities (PPFD) during the 1979–1980 ice-freeseasons. The results of these experiments could be fitted witha hyperbolic function in the manner of the Michaelis-Mentenequation (excluding rates of dark assimilation) up to –40%of the surface PPFD after which photon inhibition occurred.The half saturation constants relative to incidence PPFD (K_LT)for assimilation ( = 1.1 E m^–2d^–2) were about twice those for ( = 0.5 E m^–2d^–1).All of the K_LT values correspond to depths in thelakerangingfrom 17–29 m(–1% of surface PPFD). Dark assimilationof both and was –50% of the assimilation at saturating PPFD implying that part ofthe immediate energy required for inorganic nitrogen assimilationmay come from intermediary metabolism. This contention was supportedfor assimilation by the results of experiments performed with specific inhibitors of non-cyclic photophosphorylationand oxidative phosphorylation. The K_LT values for the assimilationof CO₂ were from 2–10 times higher than those for inorganicnitrogen. These values for CO₂ assimilation were not significantlyaltered by the addition of either or during 12 h incubations.     

Field experiments for in situ measurement of photosynthetic efficiency and quantum yield

A method for the in situ determination of     

Factors regulating summer phytoplankton in a highly eutrophic Antarctic lake

Lakes from Maritime Antarctica are regarded as systems generally inhabited by metazoan plankton capable of imposing a top-down control on the phytoplankton during short periods, while lakes from Continental Antarctica lacking these communities would be typically controlled by scarcity of nutrients, following a bottom-up model. Otero Lake is a highly eutrophic small lake located on the NW of the Antarctic Peninsula, which has no metazoan plankton. During summer 1996, we studied the density, composition and vertical distribution of the phytoplankton community of this lake with respect to various abiotic variables, yet our results demonstrated neither light nor nutrient limitation of the phytoplankton biomass. Densities of heterotrophic nanoflagellates (HNAN) and ciliates from three different size categories were also studied. Extremely low densities of HNAN (0–155 ind. ml^–1) could be due to feeding competition by bacterivore nanociliates and/or predation by large ciliates. A summer bloom of the phytoflagellate Chlamydomonas aff. celerrima Pascher reached densities tenfold those of previous years (158.10³ ind. ml^–1), though apparently curtailed by a strong peak of large ciliates (107 ind. ml^–1) which would heavily graze on PNAN (phototrophic nanoflagellates). Top-down control can thus occur in this lake during short periods of long hydrologic residence time.     

In situ stimulation of bacterial sulfate reduction in sulfate-limited freshwater lake sediments

Abstract By adding sulfate in the form of solid gypsum, it was possible to transform in situ a predominantly methanogenic sediment ecosystem into a sulfate-reducing one. The concentrations of sulfate, sulfide, methane, acetate, propionate, soluble iron, and manganese were determined in the porewater before and after the transition. Although sulfate was no longer limiting, acetate and propionate continued to accumulate and reached much higher concentrations than under sulfate-limited conditions. Metabolic activities of fermenting bacteria and of sulfate reducers, which belong to the group that incompletely oxidizes organic material, might be responsible for the increased production of volatile fatty acids. The elevated concentrations of soluble Fe(II)²⁺ and Mn(II)²⁺ observed in the porewater stem from iron and manganese compounds which may be reduced chemically by hydrogen sulfide and other microbially produced reducing agents or directly through increased activities of the iron and manganese reducing bacteria. In the horizon with high sulfate-reducing activities the methane concentrations in the porewater were lower than in non-stimulated sediment regions. The shape of the concentration depth profile indicates methane consumption through sulfate reducing processes. The in situ experiment demonstrates the response of a natural microbial ecosystem to fluctuations in the environmental conditions.     

Sedimentation of phytoplankton populations dominated by Microcystis in a shallow lake

The sedimentary flux of phytoplankton was measured using sedimenttraps in a shallow hypertrophic lake (Lake Kasumigaura), whereMicrocystis bloomed, from June to November 1983 The sedimenttraps were set at 0.5, 1.5 and 3.0 m depth in Takahamairi Bay(3.5 m depth). Microcystis spp. (including M.aerugmosa and M.viridis)in the traps were rare until early August, but increased thereafter.Sinking rates of Microcystis were 0.0045, 0.020 and 0.24 m day^–1in June–August, September and October respectively, whichwere far lower than those of Melosira (0.2–1.7 m day^–1)and Syncdra (0.2–1.0 m day^–1). The total sedimentaryfluxes of POC and that of algal carbon during the study periodwere 283.2 and 96.7 gC m^–2 which were 59.5% and 20.3%of the gross primary production (475.8 gC m^–2) respectively.The sedimentary flux of living algae measured by algal countswas large in June but small in August and September. On theother hand, the flux of detritus obtained by subtracting totalalgal carbon from POC was small in June and July but large inAugust and September. Therefore diatoms, which appeared mostlyin June, tended to sink as live algae, while Microcystis sankas detritus after being decomposed or consumed in the waterIt was concluded from the results of carbon budget calculationsand the respiration rate of the 1- to 20-µm fraction thatthe activity of decomposers or consumers increased greatly inthe short period at the end of the bloom of Microcystis.