Protistan bacterivory in an oligomesotrophic lake: Importance of attached ciliates and flagellates

Seasonal and depth variations of the abundance, biomass, and bacterivory of protozoa (heterotrophic and mixotrophic flagellates and ciliates) were determined during thermal stratification in an oligomesotrophic lake (Lake Pavin, France). Maximal densities of heterotrophic flagellates (1.9 × 10³ cells ml^–1) and ciliates (6.1 cells ml^–1) were found in the metalimnion. Pigmented flagellates dominated the flagellate biomass in the euphotic zone. Community composition of ciliated protists varied greatly with depth, and both the abundance and biomass of ciliates was dominated by oligotrichs. Heterotrophic flagellates dominated grazing, accounting for 84% of total protistan bacterivory. Maximal grazing impact of heterotrophic flagellates was 18.9 × 10⁶ bacteria 1^–1h^–1. On average, 62% of nonpigmented flagellates were found to ingest particles. Ciliates and mixotrophic flagellates averaged 13% and 3% of protistan bacterivory, respectively. Attached protozoa (ciliates and flagellates) were found to colonize the diatom Asterionella formosa. Attached bacterivores had higher ingestion rates than free bacterivorous protozoa and may account for 66% of total protozoa bacterivory. Our results indicated that even in low numbers, epibiotic protozoa may have a major grazing impact on free bacteria. Correspondence: C. Amblard.     

Biomass production in mixotrophic culture of Euglena gracilis under acidic condition and its growth energetics

When Euglena gracilis was grown in the heterotrophic condition with glucose and (NH₄)₂SO₄ as the carbon and nitrogen source, a high cell yield (4.28–4.48 g l^–1) was obtained and the culture pH decreased to 1.6–2. The biomass production in the heterotrophic culture was compared to those in the autotrophic and mixotrophic cultures. Autotrophic growth was 4.7–6.3% of the heterotrophic one, whereas about 15–19% higher growth was obtained in the mixotrophic culture. Moreover, good production of chlorophyll (39.4 mg l^–1) and carotenoids (13.8 mg l^–1) were attained in the mixotrophic culture, giving the highest fermenter productivity with respect to biomass as well as chlorophyll and carotenoids. Through an energetic analysis in the mixotrophic culture, it was estimated about 25–28% of the total ATP requirement is formed in the photochemical reactions. This resulted in an improved biomass production in the mixotrophic culture of E. gracilis.     

Seasonal cycle of the microbial plankton in Crooked Lake,Antarctica

Summary Changes in the abundance of the components of the microbial plankton between July 1990 and March 1991 in Crooked Lake, one of the largest and deepest freshwater lakes in Antarctica, are described. Chlorophyll a concentration is low (0.2–0.4g·1^–1) and there is no discernable spring increase. The phytoplankton is largely dominated by flagellates. Bacterioplankton exhibits a seasonal pattern of abundance ranging from 1.0 × 10⁸·1^–1 in July to 3.25 × 10⁸·1^–1 in September. Changes in bacterial abundance probably relate to temperature and grazing by heterotrophic and mixotrophic flagellates. Total flagellated protozoan concentrations ranged between 25–136 × 10²·l^–1. Autotrophic and heterotrophic flagellate abundances were coupled and peaks in their abundance oscillated with peaks in bacterioplankton concentration. Four species of ciliated protozoa, dominated by oligotrichs, particularly the plastidic Strombidium, inhabit the lake. The plankton is characterised by the presence of floes which act as loci for bacteria, flagellates and amoebae and feeding sites for the ciliates and the two sparce metazoan components of the plankton. Crooked Lake is extremely oligotrophic but nonetheless supports a plankton community with a low species diversity and simple trophodynamics.     

Phytoplankton, bacterial production and protozoan bacterivory in stratified, brackish-water Lake Shira (Khakasia, Siberia)

Rates of oxygenic and anoxygenic photosynthesis, chemoautotrophic and heterotrophic bacterial production and protozoan bacterivory were measured in the pelagic zone of the stratified brackish-water lake with the purpose to determine the vertical distribution of these processes and to estimate their significance in the functioning of planktonic community of the lake. In midsummer, total daily primary productivity was about 1.3 g C m^–2, of which 72% was produced by the phytoplankton, 24% by the chemoautotrophic bacteria, and only 4% by the phototrophic sulphur bacteria. Thus anoxygenic photosynthesis is a negligible source of organic matter in the lake. The production of heterotrophic bacteria averaged 1.5 g C m^–2 d^–1 and exceeded the total photosynthesis of phytoplankton and photosynthetic bacteria by a factor of 1.5. The estimated total primary production was too low to sustain the bacterial production. Probably the carbon cycle in the lake is dependent on the input of allochthonous organic matter. As a rule, the maximal rates of primary production and heterotrophic bacterial production were found in the chemocline or at the upper boundary of the chemocline. Heterotrophic flagellates dominated among the protozoan populations and were the major consumers of the bacterioplankton production in the lake. They showed maximal ingestion rates from 2.3 to 2.9 mg C m^–3 h^–1 at the upper boundary of the chemocline, where they consumed from 50 to 54% of the production of heterotrophic bacteria. Data obtained indicate that in Lake Shira the oxic-anoxic interface is the site of the most intensive production and mineralization of organic matter.     

Structure of planktonic microbial food web in a brackish stratified Siberian lake

The distribution of primary components of the microbial community (autotrophic pico- and nanoplankton, phototrophic bacteria, heterotrophic bacteria, microscopic fungi, heterotrophic flagellates, ciliates and heliozoa) in the water column of Lake Shira, a steppe brackish-water, stratified lake in Khakasia, Siberia (Russia), were assessed in midsummer. Bacterioplankton was the main component of the planktonic microbial community, accounting for 65.3 to 75.7% of the total microbial biomass. The maximum concentration of heterotrophic bacteria were recorded in the monimolimnion of the lake. Autotrophic microorganisms contributed more significantly to the total microbial biomass in the pelagic zone (20.2–26.5%) than in the littoral zone of the lake (8.7–14.9%). First of all, it is caused by development of phototrophic sulphur bacteria at the oxic-anoxic boundary. The concentrations of most aerobic phototrophic and heterotrophic microorganisms were maximal in the upper mixolimnion. Heterotrophic flagellates dominated the protozoan populations. Ciliates were minor component of the planktonic microbial community of the lake. Heterotrophic flagellates were the most diverse group of planktonic eucaryotes in the lake, which represented by 36 species. Facultative and obligate anaerobic flagellates were revealed in the monimolimnion. There were four species of Heliozoa and only three of ciliates in the lake.     

Occurrence of mixotrophic flagellates in relation to bacterioplankton production, light regime and availability of inorganic nutrients in unproductive lakes with differing humic contents

1. Field data from five unproductive Swedish lakes were used to investigate the occurrence of mixotrophic flagellates in relation to bacterioplankton, autotrophic phytoplankton, heterotrophic flagellates and abiotic environmental factors. Three different sources of data were used: (i) a 3‐year study (1995–97) of the humic Lake Örträsket, (ii) seasonal measurements from five lakes with widely varying dissolved organic carbon (DOC) concentrations, and (iii) whole lake enrichment experiments with inorganic nutrients and organic carbon. 2. Mixotrophic flagellates usually dominated over autotrophic phytoplankton in Lake Örträsket in early summer, when both bacterial production and light levels were high. Comparative data from the five lakes demonstrated that the ratio between the biomasses of mixotrophic flagellates and autotrophic phytoplankton (the M/A‐ratio) was positively correlated to bacterioplankton production, but not to the light regime. Whole lake carbon addition (white sugar) increased bacterial biomass, and production, reduced the biomass of autotrophs by a factor of 16, and increased the M/A‐ratio from 0.03 to 3.4. Collectively, the results indicate that the dominance of mixotrophs among phytoplankton was positively related to bacterioplankton production. 3. Whole lake fertilisation with nitrogen (N) and phosphorus (P) demonstrated that the obligate autotrophic phytoplankton was limited by N. N‐addition increased the biomass of the autotrophic phytoplankton but had no effect on mixotrophic flagellates or bacteria, and the M/A‐ratio decreased from 1.2 to 0.6 after N‐enrichment. Therefore, we suggest that bacteria under natural conditions, by utilising allochthonous DOC as an energy and carbon source, are able to outcompete autotrophs for available inorganic nutrients. Consequently, mixotrophic flagellates can become the dominant phytoplankters when phagotrophy permits them to use nutrients stored in bacterial biomass. 4. In Lake Örträsket, the biomass of mixotrophs was usually higher than the biomass of heterotrophs during the summer. This dominance could not be explained by higher grazing rates among the mixotrophs. Instead, ratios between mixotrophic and heterotrophic biomass (the M/H‐ratio) were positively related to light availability. Therefore, we suggest that photosynthesis can enable mixotrophic flagellates to outcompete heterotrophic flagellates.     

Production potential of eicosapentaenoic acid by the diatom Nitzschia laevis

To investigate the production potential of eicosapentaenoic acid (EPA) by the diatom Nitzschia laevis, the growth characteristics and fatty acid composition of the cells were studied under photoautotrophic, mixotrophic and heterotrophic conditions of growth. The specific growth rate and maximum biomass concentration were respectively 0.466 d^–1 and 2.27 g l^–1 for mixotrophic culture, 0.344 d^–1 and 2.04 g l^–1 for heterotrophic culture, and 0.167 d^–1 and 0.5 g l^–1 for photoautotrophic culture, respectively. As for EPA production, the yield and productivity were respectively 52.32 mg l^–1 and 10.46 mg l^–1 d for mixotrophic culture, 35.08 mg l^–1 and 6.37 mg l^–1 d for heterotrophic culture, and 6.78 mg l^–1 and 3.39 mg l^–1 d for photoautotrophic culture, respectively. Results suggest that mixotrophic culture is the most suitable growth mode for the production of EPA by the diatom Nitzschia laevis. The results are useful for the development of a cost-effective fermentation process for EPA production by Nitzschia laevis.     

Microbial Community Structure and Dynamics in the Largest Natural French Lake (Lake Bourget)

We investigated the dynamics and diversity of heterotrophic bacteria, autotrophic and heterotrophic flagellates, and ciliates from March to July 2002 in the surface waters (0–50 m) of Lake Bourget. The heterotrophic bacteria consisted mainly of “small” cocci, but filaments (>2 μm), commonly considered to be grazing-resistant forms under increased nanoflagellate grazing, were also detected. These elongated cells mainly belonged to the Cytophaga-Flavobacterium (CF) cluster, and were most abundant during spring and early summer, when mixotrophic or heterotrophic flagellates were the main bacterial predators. The CF group strongly dominated fluorescent in situ hybridization–detected cells from March to June, whereas clear changes were observed in early summer when Beta-proteobacteria and Alpha-proteobacteria increased concomitantly with maximal protist grazing pressures. The analysis of protist community structure revealed that the flagellates consisted mainly of cryptomonad forms. The dynamics of Cryptomonas sp. and Dinobryon sp. suggested the potential importance of mixotrophs as consumers of bacteria. This point was verified by an experimental approach based on fluorescent microbeads to assess the potential grazing impact of all protist taxa in the epilimnion. From the results, three distinct periods in the functioning of the epilimnetic microbial loop were identified. In early spring, mixotrophic and heterotrophic flagellates constituted the main bacterivores, and were regulated by the availability of their resources mainly during April (phase 1). Once the “clear water phase” was established, the predation pressure of metazooplankton represented a strong top-down force on all microbial compartments. During this period only mixotrophic flagellates occasionally exerted a significant bacterivory pressure (phase 2). Finally, the early summer was characterized by the highest protozoan grazing impact and by a rapid shift in the carbon pathway transfer, with a fast change-over of the main predators contribution, i.e., mixotrophic, heterotrophic flagellates and ciliates in bacterial mortality. The high abundance of ciliates during this period was consistent with the high densities of resources (heterotrophic nanoflagellates, algae, bacteria) in deep layers containing the most chlorophyll. Bacteria, as ciliates, responded clearly to increasing phytoplankton abundance, and although bacterial grazing impact could vary largely, bacterial abundance seemed to be primarily bottom-up regulated (phase 3).     

Zooplankton-mediated changes of bacterial community structure

Enclosure experiments in the mesotrophic Schöhsee in northern Germany were designed to study the impact of metazooplankton on components of the microbial food web (bacteria, flagellates, ciliates). Zooplankton was manipulated in 500-liter epilimnetic mesocosms so that either Daphnia or copepods were dominating, or metazooplankton was virtually absent. The bacterial community responded immediately to changes in zooplankton composition. Biomass, productivity, and especially the morphology of the bacteria changed drastically in the different treatments. Cascading predation effects on the bacterioplankton were transmitted mainly by phagotrophic protozoans which had changed in species composition and biomass. When Daphnia dominated, protozoans were largely suppressed and the original morphological structure of the bacteria (mainly small rods and cocci) remained throughout the experiment. Dominance of copepods or the absence of metazoan predators resulted in a mass appearance of bacterivorous protists (flagellates and ciliates). They promoted a fast decline of bacterial abundance and a shift to the predominance of morphologically inedible forms, mainly long filaments. After 3 days they formed 80–90% of the bacterial biomass. The results indicate that metazooplankton predation on phagotrophic protozoans is a key mechanism for the regulation of bacterioplankton density and community structure.Correspondence to: K. Jürgens.     

The Soil Flagellate Heteromita globosa Accelerates Bacterial Degradation of Alkylbenzenes through Grazing and Acetate Excretion in Batch Culture

The impact of grazing by soil flagellates Heteromita globosa on aerobic biodegradation of benzene by Pseudomonas strain PS+ was examined in batch culture. Growth of H. globosa on these bacteria obeyed Monod kinetics (_max, 0.17 ± 0.03 h^–1; K_s, 1.1 ± 0.2 × 10⁷ bacteria mL^–1) and was optimal at a bacteria/ flagellate ratio of 2000. Carbon mass balance showed that 5.2% of total [ring-U-¹⁴C]benzene fed to bacteria was subsequently incorporated into flagellate biomass. Growth-inhibiting concentrations (IC₅₀) of alkylbenzenes (benzene, toluene, ethylbenzene) were inversely related with their octanol/ water partitioning coefficients, and benzene was least toxic for bacteria and flagellates with IC₅₀ values of 4392 (± 167) M and 2770 (± 653) M, respectively. The first-order rate constant for benzene degradation (k₁, 0.48 ± 0.12 day^–1) was unaffected by the presence or absence of flagellates in cultures. However, the rate of benzene degradation by individual bacteria averaged three times higher in the presence of flagellates (0.73 ± 0.13 fmol cell^–1 h^–1) than in their absence (0.26 ± 0.03 fmol cell^–1 h^–1). Benzene degradation also coincided with higher levels of dissolved oxygen and a higher rate of nitrate reduction in the presence of flagellates (p < 0.02). Grazing by flagellates may have increased the availability of dissolved oxygen to a smaller surviving population of bacteria engaged in the aerobic reactions initiating benzene degradation. In addition, flagellates may also have increased the rate of nitrate reduction through the excretion of acetate as an additional electron donor for these bacteria. Indeed, acetate was shown to progressively accumulate in cultures where flagellates grazed on heat-killed bacteria. This study provided evidence that grazing flagellates stimulate bacterial degradation of alkylbenzenes and provide a link for carbon cycling to consumers at higher trophic levels. This may have important implications for bioremediation processes.     

Limnology of Crater Lakes in Los Tuxtlas,Mexico

Investigations on the abundance, biomass and position of heterotrophic flagellates (HF) in the benthic microbial food web of a melt water stream on King George Island, Antarctic Peninsula, were undertaken during the Antarctic summer from 23rd December 1997 until 13th March 1998. Abundance and biomass of potential HF resources (picophotoautotrophic and non-photoautotrophic bacteria) as well as potential predators on HF (ciliates and meiofauna) were also investigated. HF abundance ranged from approximately 9 × 10³ to 81 × 10³ cells cm^–3, values which fall into the same range as those found in lower latitudes. Numerically important benthic HF were euglenids, kinetoplastids, thaumatomastigids and especially chrysomonads. Most species identified have been shown to have a worldwide distribution. Abundance of the benthic ciliates ranged from 27 to 950 cells cm^–3. Mean bacterial abundance was 1.9 × 10⁷ and 5.2 × 10⁸ cells cm^–3 for picophotoautotrophic and non-photoautotrophic benthos, respectively. The well-developed microbial community was able to support the large number of nematods, gastotrichs, tardigrads and rotifers with abundances reaching more than 1000 individuals cm^–3. The largest portion of heterotrophic biomass was formed by the meiofauna with a mean of 63 g C cm^–3, followed by that of the heterotrophic bacteria with 4.80 g C cm^–3. Picophotoautotrophic bacteria contributed a mean of 1.37 g C cm^–3. HF and ciliates mean biomass was 0.61 and 1.99 g C cm^–3, respectively, with the HF biomass comprising between <10 and 70% of the total protozoan biomass. The data obtained in this study identify the melt water stream as a hot-spot of heterotrophic microbial and meiofaunal activity during the austral summer. The HF in the melt water stream formed a diverse group in terms of taxa and potential feeding types. Chrysomonads, kinetoplastids, euglenids and thaumatomastigida were the most abundant taxa. A classification into feeding types identified an average of 34% of the total HF as bacterivorous while all others were able to utilise other, larger organisms as resources. Potential trophic interactions between HF and bacteria and higher trophic levels are discussed.     

Abundance and biomass of heterotrophic microorganisms in Lake Tanganyika

1. This study focused on heterotrophic microorganisms in the two main basins (north and south) of Lake Tanganyika during dry and wet seasons in 2002. Bacteria (81% cocci) were abundant (2.28–5.30 × 10⁶ cells mL^?1). During the dry season, in the south basin, bacterial biomass reached a maximum of 2.27 g C m^?2 and phytoplankton biomass was 3.75 g C m^?2 (integrated over a water column of 100 m). 2. Protozoan abundance was constituted of 99% of heterotrophic nanoflagellates (HNF). Communities of flagellates and bacteria consisted of very small but numerous cells. Flagellates were often the main planktonic compartment, with a biomass of 3.42–4.43 g C m^?2. Flagellate biomass was in the same range and often higher than the total autotrophic biomass (1.60–4.72 g C m^?2). 3. Total autotrophic carbon was partly sustained by the endosymbiotic zoochlorellae Strombidium. These ciliates were present only in the euphotic zone and usually contributed most of the biomass of ciliates. 4. Total heterotrophic ciliate biomass ranged between 0.35 and 0.44 g C m^?2. In 2002, heterotrophic microorganisms consisting of bacteria, flagellates and ciliates represented a large fraction of plankton. These results support the hypothesis that the microbial food web contributes to the high productivity of Lake Tanganyika. 5. As the sole source of carbon in the pelagic zone of this large lake is phytoplankton production, planktonic heterotrophs ultimately depend on autochthonous organic carbon, most probably dissolved organic carbon (DOC) from algal excretion.     

Plankton community of a polyhumic lake with and without Daphnia longispina (Cladocera)

The impact of Daphnia longispina (Cladocera) on the plankton food web was studied in a polyhumic lake where this species comprised almost all zooplankton biomass. Plastic enclosures (volume 7 m³) were inserted into the lake retaining the initial water stratification except that in one enclosure zooplankton was removed. After the removal of Daphniaa rotifer, Keratella cochlearis, ciliates and heterotrophic nanoflagellates increased markedly and the density and biomass of bacteria decreased. Edible algal species, Cryptomonas rostratiformisand three small chrysophytes,Ochromonas, Pedinella and Spinifermonas, took advantage of the removal of Daphnia, while more grazing-resistant species declined. In spite of the changes in the species composition of phytoplankton, the removal of Daphnia did not affect the biomass, primary production or respiration of plankton. The results implied that the density of heterotrophic flagellates and ciliates was controlled by Daphnia, but in its absence the former took its role as the bacterial grazers.     

Development of ice biota in a temperate sea area (Gulf of Bothnia)

A study of sea ice biota was carried out in the Gulf of Bothnia (northern Baltic Sea) during the winter of 1989–1990. Samples (ice cores) were taken at a coastal station at regular time intervals during the ice season. Chlorophyll a concentration, algal species distribution, bacterial numbers, and primary and bacterial production were measured. Colonization of the ice began in January when daylight was low. As the available light increased, the algae started to grow exponentially. The vertical chlorophyll a distribution changed and algal species composition and biomass changed during the season. During the initial and middle phase of colonization, ice-specific diatoms, Nitzschia frigida and Navicula pelagica, dominated the algal biomass. Nutrients (PO₄ ^3– and NO_3j) were found to be depleted during the time of algal exponential growth. The maximum algal biomass exceeded 800 g C 1^–1. The primary production supplied food for heterotrophic organisms. The presence of heterotrophic organisms of different trophic levels (bacteria, flagellates, ciliates and rotifers) indicated an active microbial food web.     

Nutritional characteristics of a mixotrophic nanoflagellate,Ochromonas sp.

Autotrophic and heterotrophic growth characteristics of a nano-flagellate were investigated. The flagellate,Ochromonas sp., was isolated from the northern Baltic Sea. Autotrophic growth was poor. Axenically pregrown flagellates did not increase significantly in cell number during incubation in different inorganic media. The number of flagellates remained constant 3–5 weeks in cultures kept in the light (100mol m^–2 sec^–1), whereas in the dark, a high mortality rate was found. Uptake of inorganic¹⁴C into an acid-stable fraction indicated thatOchromonas had a functional photosynthetic apparatus. Heterotrophic growth in both liquid medium and medium containing bacteria was rapid. The maximum growth rate corresponded to a generation time of 5.3 hours. Light had no effect on heterotrophic growth. Cells pregrown onEscherichia coli minicells survived without additional bacteria as food when kept in the light, but rapid death occurred in darkness. In conclusion, heterotrophy is the major mechanism to support growth in this species ofOchromonas, but under poor environmental conditions photoautotrophy might be a strategy for survival rather than growth.     

Patterns in pelagic and benthic nanoflagellate densities in the coastal upwelling system along the Banc d'Arguin,Mauritania

We studied spatial variation in abundance of marine benthic and pelagic heterotrophic nanoflagellates in relation to abundances of autotrophic flagellates, bacteria and cyanobacteria in an upwelling area off the Banc d'Arguin, Mauritania.There was enormous spatial variation in densities. In the sediments these ranged from 8–219 × 10³ cm^–3 for heterotrophic flagellates. Maximum values are in the range of those for temperate shallow marine bottoms. Low densities (< 20 × 10³) were confined to the deep stations (> 1000 m). Over the shelf (10–100 m depth) densities were high but related to grain size rather than to concurrent upwelling phenomena or to the abundance of benthic macrofauna.Pelagic flagellate abundance appeared to be more indicative of contemporary hydrographic conditions, most obvious by an increase in the ratio heterotrophic/autotrophic nanoflagellates away from the area of most intense upwelling.     

Vertical distribution in and isolation of bacteria from Lake Vanda: an Antarctic lake

Vertical distribution of bacteria in Lake Vanda, an Antarctic meromictic lake, was examined by the acridine orange epifluorescence direct count method. Total bacteria were 10⁴–10⁵ cells · ml^–1 in the water at 55 m depth and above, and increased drastically to 10⁷ cells · ml^–1 in the bottom water. Filamentous or long rodshaped bacteria occurred at a high frequency in the upper layers, but in the bottom layers most bacteria were coccoidal or short rods. Mean bacterial cell volume in water of between 10 m and 60 m deep was fairly large compared with common bacterial populations in seawater and lake water. Aerobic heterotrophic bacteria were recovered from the water of a depth of 30 m and above, and were assumed to belong to Caulobacter. Viable heterotrophic bacteria were not recovered from the high salinity deep water by media prepared with the same deep water. Phototrophic purple non-sulphur bacteria were isolated by enrichment cultures from water at 55 m depth.     

Growth characteristics of the cyanobacterium Nostoc flagelliforme in photoautotrophic, mixotrophic and heterotrophic cultivation

Nostoc flagelliforme is a terrestrial cyanobacterium with high economic value. Dissociated cells separated from a natural colony of N. flagelliforme were cultivated for 7 days under either phototrophic, mixotrophic or heterotrophic culture conditions. The highest biomass, 1.67 g L⁻¹ cell concentration, was obtained under mixotrophic culture, representing 4.98 and 2.28 times the biomass obtained in phototrophic and heterotrophic cultures, respectively. The biomass in mixotrophic culture was not the sum as that in photoautotrophic and heterotrophic cultures. During the first 4 days of culture, the cell concentration in mixotrophic culture was lower than the sum of those in photoautotrophic and heterotrophic cultures. However, from the 5th day, the cell concentration in mixotrophic culture surpassed the sum of those obtained from the other two trophic modes. Although the inhibitor of photosynthetic electron transport DCMU [3-(3,4-dichlorophenyl)-1,1-dimethylurea] efficiently inhibited autotrophic growth of N. flagelliforme cells, under mixotrophic culture they could grow by using glucose. The addition of glucose changed the response of N.flagelliforme cells to light. The maximal photosynthetic rate, dark respiration rate and light compensation point in mixotrophic culture were higher than those in photoautotrophic cultures. These results suggest that photoautotrophic (photosynthesis) and heterotrophic (oxidative metabolism of glucose) growth interact in mixotrophic growth of N. flagelliforme cells.     

Interactions between metazoans,autotrophs, mixotrophs and bacterioplankton in nutrient‐depleted high DOC environments: a long‐term experiment

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Planktonic food chains of a highly humic lake

The development and metabolism of the plankton of a highly humic lake were followed over the vernal primary production maximum. The study was made in a mesocosm in which large filter feeders, typical of this lake in summer, were absent. During the rising phase of phytoplankton, the community was predominantly autotrophic. The most important constituents in the algal biomass were a dinoflagellate, Gymnodinium sp. (40–50%), and a prasinophycean, Scourfieldia cordiformis (7%). The biomasses of Chlamydomonas spp. and Chrysococcus spp. reached their maxima a few days later and Cryptomonas sp. became most abundant at the end of the experiment. After the phytoplankton maximum, about one week from the beginning ofthe experiment, grazing of algae by phagotrophic protozoans and phosphate depletion led to a rapid decrease of algal biomass and the community became predominantly heterotrophic. In spite of a large variation in algal biomass and primary production, the biomass of bacteria remained of the same order of magnitude as in algae both before and after the algal maximum. Bacteria were mostly responsible for the plankton respiration, which also showed no dependence on primary production. Since exudation by phytoplankton was also low, the nutrition of bacterioplankton was probably mainly based on allochthonous dissolved organic matter rather than or primary production. Thus the production of bacteria was an additional food source for higher trophic levels along with phytoplankton. Because filter feeding zooplankton was absent in the experiment, protozoans were the only grazers utilizing algae and bacteria. Essentially all growth of bacteria was used by bacterivores.