Bacterioplankton Growth on Fractions of Dissolved Organic Carbon of Different Molecular Weights from Humic and Clear Waters

The ability of fractions of dissolved organic carbon (DOC) of different molecular weights (MW) to support bacterial growth was studied in batch culture experiments. Natural pelagic bacteria were inoculated into particle-free (0.2-μm filtered) water, taken from 10 oligotrophic lakes of differing humic content, and either used without further treatments or ultrafiltered to remove DOC of >10,000 MW or >1,000 MW. Stationaryphase abundance of bacteria in the cultures was used as an estimate of bacterial carrying capacity. High-MW DOC (>10,000) comprised an increasing fraction of total DOC with increasing total DOC and humic content of the lakes. High-MW DOC was generally more available to bacteria (i.e., more bacteria were produced per unit of organic carbon initially present) than low-MW (<10,000) DOC. The availability to bacteria of this high-MW DOC decreased with increasing humic content. However, although less available in humic lakes than in clearwater lakes, the higher abundance of high-MW DOC made it quantitatively more important as a bacterial substrate; i.e., a larger fraction of the total bacterial yield of the cultures was due to high-MW DOC compounds in humic lakes than in clearwater lakes. On the average, 48% of bacterial growth occurred at the expense of DOC of <10,000 MW. DOC of <1,000 MW was responsible for an average of 22% of bacterial growth, with no significant correlation to humic content and DOC concentration of lakes. The DOC which supports bacterial growth, as well as the total DOC, is of different quality in humic and clearwater lakes.     

Effects of Additions of DOC on Pelagic Biota in a Clearwater System: Results from a Whole Lake Experiment in Northern Sweden

An oligotrophic clearwater lake, initially characterized by a pronounced dominance of autotrophic phytoplankton and mostly by one species, the green alga Botryococcus, was subject to additions of dissolved organic carbon in the form of white sugar (sucrose) during two consecutive years. The hypothesis tested was that it is organic carbon per se, and not other possible effects of humic substances, that determines the differences in structure of the planktonic ecosystem between humic and clearwater lakes. The additions of DOC resulted in a significant increase in bacterial biomass and a decrease in the biomass of autotrophic phytoplankton. The biomass of mixotrophic and heterotrophic flagellates instead increased significantly, whereas no effects were found to propagate to higher trophic levels. As a result of the changes among biota, total planktonic biomass also decreased to a level typical of nearby humic lakes. We suggest that it is the carbon component of humic material and its utilization by bacterioplankton that determines the structure and function of the pelagic food web in humic lakes.     

Bacterial Growth in Mixed Cultures on Dissolved Organic Carbon from Humic and Clear Waters

Interactions between bacterial assemblages and dissolved organic carbon (DOC) from different sources were investigated. Mixed batch cultures were set up with water from a humic and a clear-water lake by a 1:20 dilution of the bacterial assemblage (1.0 μm of prefiltered lake water) with natural medium (sterile filtered lake water) in all four possible combinations of the two waters and their bacterial assemblages. Bacterial numbers and biomass, DOC, thymidine incorporation, ATP, and uptake of glucose and phenol were followed in these cultures. Growth curves and exponential growth rates were similar in all cultures, regardless of inoculum or medium. However, bacterial biomass produced was double in cultures based on water from the humic lake. The fraction of DOC consumed by heterotrophic bacteria during growth was in the same range, 15 to 22% of the total DOC pool, in all cultures. Bacterial growth efficiency, calculated from bacterial biomass produced and DOC consumed, was in the order of 20%. Glucose uptake reached a peak during exponential growth in all cultures. Phenol uptake was insignificant in the cultures based on the clear-water medium, but occurred in humic medium cultures after exponential growth. The similarity in the carbon budgets of all cultures indicated that the source of the bacterial assemblage did not have a significant effect on the overall carbon flux. However, fluxes of specific organic compounds differed, as reflected by glucose and phenol uptake, depending on the nature of the DOC and the bacterial assemblage.     

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

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

Viral and Bacterioplankton Dynamics in Two Lakes with Different Humic Contents

Viral and bacterioplankton dynamics were investigated, together with the temporal variation of phage-infected bacterioplankton in two oligotrophic lakes, one humic and the other clearwater. Bacterial abundance was significantly higher in the humic lake, while the abundance of virus-like particles (VLP) was significantly higher in the clearwater lake. There were no differences in either the frequency of infected bacterial cells (FIC), or in burst size between the lakes. Because of the higher bacterial abundance in the humic lake, a larger number of bacteria were lyzed in this lake. FIC showed large seasonal changes, varying between 9 and 43%, which covers almost the entire range of previously published data from both lacustrine and marine environments. The temporal changes in VLP abundance and FIC were slow in both the humic and clearwater lakes. The burst size was low in both lakes (average value, nine in each case), probably because of the oligotrophic status of the lakes. The chlorophyll a concentrations were higher and positively correlated with VLP numbers in the clearwater lake, indicating that a significant proportion of the viruses in this lake may be phytoplankton viruses.     

Colloidal and dissolved organic matter in lake water: Carbohydrate and amino acid composition,and ability to support bacterial growth

Bacterial utilization of dissolved organic matter (DOM) was studied in water from a humic and a clearwater oligotrophic lake. Indigenous bacteria were inoculated into either 0.2 m natural filtered lake water, or lake water enriched fivefold with colloidal DOM >100 kD but below 0.2 m. Consumption of DOM was followed from changes in concentrations of total dissolved organic carbon (DOC), dissolved combined and free carbohydrates and amino acids (DCCHO and DFCHO, and DCAA and DFAA, respectively) and by uptake of monosaccharide and amino acid radioisotopes. DCCHO and DCAA made up 8% (humic lake) to 33–44% (clear-water lake) of the natural DOC pools, while DFCHO and DFAA contributed at most 1.7% to the DOC pools. Addition of >100 kD DOM increased the DOC concentrations by 50% (clearwater lake) to 92% (humic lake), but it only resulted in a higher bacterial production (by 63%) in the humic lake. During the incubations 13 to 37% of the DOC was assimilated by the bacteria, at estimated growth efficiencies of 4–8%. Despite the measured reduction of DOC, statistically significant changes of specific organic compounds, especially of DCCHO and DCAA, generally did not occur. Probably the presence of high molecular weight DOC interfered with the applied analytical procedures. Addition of radiotracers indicated, however, that DFAA sustained 17–58% and 29–100% of the bacterial carbon and nitrogen requirements, respectively, and that glucose met 1–3% of the bacterial carbon requirements. Thus, our experiments indicate that radiotracers, rather than measurements of concentration changes, should be used in studies of bacterial utilization of DOC in freshwaters with a high content of humic or high molecular weight organic matter.     

Distribution of planktonic ciliates in highly coloured subtropical lakes: comparison with clearwater ciliate communities and the contribution of myxotrophic taxa to total autotrophic biomass

SUMMARY 1. The planktonic ciliate communities of eleven organically coloured north and central Florida lakes representing a variety of trophic conditions were examined during 1979–80. The total abundance and biomass of ciliates were not significantly different from comparable clearwater lakes and only minor taxonomic replacements were noted at the order level.
2. Timing of population peaks of oligotrophic lakes was dissimilar to clearwater lakes of the same trophic state, but seasonality in meso-trophic and eutrophic lakes resembled patterns described for comparable clearwater lakes.
3. Various ciliate components were strongly correlated with chlorophyll a concentrations, but only moderately correlated to dominant phytoplankton groups. No significant correlations were found between ciliate components and bacterial abundance.
4. Myxotrophic taxa numerically dominated oligotrophic systems, particularly during midsummer, and accounted for a large percentage of the total ciliate biomass. Estimates of the ciliate contribution to total autotrophic biomass indicate that these zoochlorellae-bearing protozoa may account for much of the autotrophic biomass during midsummer periods in coloured lakes, and thus may lead to an overestimation of phytoplankton standing crops available to zooplankton grazers if chlorophyll a is used as a surrogate measure of algal biomass.     

Bacterioplankton growth, grazing mortality and quantitative relationship to primary production in a humic and a clearwater lake

Bacterial growth and grazing mortality were estimated from Mayto October in two south Swedish oligotrophic lakes, one beinga clearwater lake (water colour 5–10 mg Pt l^–1 DOC2.9–3.4 mg l^–1, Secchi disk depth 5.0–9.4m) and the other a humic, brownwater lake (water colour 105–165mg Pt l^–1, DOC 13.7–22.7mg l^–1, Secchi diskdepth 1.3–2.1 m). Specific rates of growth and grazingmortality were generally similar for both lakes. However, theabundance of bacteria was consistently 2–3 times higherin the water of the humic lake, suggesting that the total productionand consumption of bacterial cells were also higher than inthe dearwater lake. The ratio of bacterial secondary productionto primary production was higher in the humic lake than in theclearwater lake, indicating that the bacterioplankton of thehumic lake utilize allochthonous substrates, in addition tosubstrates originating from autochthonous primary production.Most of the bacterial loss in both lakes could be attributedto small protozoan grazers. This implies that allochthonousand autochthonous organic carbon fixed by bacterioplankton isless important in terms of carbon flow to higher trophic levelsthan would be expected if macrozooplankton were the dominantbacterivores, providing a more direct and efficient transferof carbon to larger organisms.     

Heterotrophic bacterial growth efficiency and community structure at different natural organic carbon concentrations

Batch cultures of aquatic bacteria and dissolved organic matter were used to examine the impact of carbon source concentration on bacterial growth, biomass, growth efficiency, and community composition. An aged concentrate of dissolved organic matter from a humic lake was diluted with organic compound-free artificial lake water to obtain concentrations of dissolved organic carbon (DOC) ranging from 0.04 to 2.53 mM. The bacterial biomass produced in the cultures increased linearly with the DOC concentration, indicating that bacterial biomass production was limited by the supply of carbon. The bacterial growth rate in the exponential growth phase exhibited a hyperbolic response to the DOC concentration, suggesting that the maximum growth rate was constrained by the substrate concentration at low DOC concentrations. Likewise, the bacterial growth efficiency calculated from the production of biomass and CO(2) increased asymptotically from 0.4 to 10.4% with increasing DOC concentration. The compositions of the microbial communities that emerged in the cultures were assessed by separation of PCR-amplified 16S rRNA fragments by denaturing gradient gel electrophoresis. Nonmetric multidimensional scaling of the gel profiles showed that there was a gradual change in the community composition along the DOC gradient; members of the beta subclass of the class Proteobacteria and members of the Cytophaga-Flavobacterium group were well represented at all concentrations, whereas members of the alpha subclass of the Proteobacteria were found exclusively at the lowest carbon concentration. The shift in community composition along the DOC gradient was similar to the patterns of growth efficiency and growth rate. The results suggest that the bacterial growth efficiencies, the rates of bacterial growth, and the compositions of bacterial communities are not constrained by substrate concentrations in most natural waters, with the possible exception of the most oligotrophic environments.     

Bacterial growth on dissolved organic carbon from a blackwater river

Different nominal molecular weight (nMW) fractions of DOC from a southeastern blackwater river were concentrated by ultrafiltration and added to sieved river water to assess each fraction's ability to stimulate bacterial growth. Bacterial growth was measured using change in bacterial biomass from direct counts and using³H-thymidine incorporated into DNA. Bacterial growth and amount of DOC used was greatest in the low MW enrichment (< 1,000 nMW) and least in the intermediate MW enrichment (1,000–10,000 nMW). The high MW fraction (> 10,000 nMW) supported more growth than did the intermediate MW fraction, apparently because of lower MW compounds complexed with a high MW refractory core. The low MW fraction of DOC from a clearwater mountain stream, a boreal blackwater river, and leachate from water oak and willow leaves also stimulated more bacterial growth than did other fractions. However, the high MW DOC from these other sources was not as biologically available as high MW DOC from a blackwater river. Bacteria converted blackwater river DOC to bacterial biomass with an efficiency of 31%. Bacteria produced at the expense of abundant riverine DOC provide a trophic resource for protozoa and higher levels of the microbial food web of a blackwater river.     

Relationships between DOC concentration and epilithon stable isotopes in boreal lakes

1. Littoral biota in boreal lakes are known to assimilate epilithon. Being able to predict the stable isotopic composition of these alga will help to identify those systems in which δ¹³C and δ¹⁵N analysis can be used in foodweb investigations of allochthony and biomagnification.
2. In a survey of 15 boreal lakes, the concentration of dissolved organic carbon (DOC) explained 76% of the variation in epilithon δ¹³C, and 86% of the variation in epilithon δ¹⁵N.
3. Because both δ¹³C and δ¹⁵N values were depressed and similar to terrestrial values in humic (high DOC) lakes, it will be more difficult to successfully employ stable isotopic techniques for estimating allochthony in such systems. Lower δ¹⁵N values in humic lakes also indicate that trophic positions estimated by stable isotopes are not directly comparable to those of similar biota inhabiting clearwater lakes, unless autochthonous baseline corrections are made.     

Effect of exposure to sunlight and phosphorus-limitation on bacterial degradation of coloured dissolved organic matter (CDOM) in freshwater

This study reports on the interacting effect of photochemical conditioning of dissolved organic matter and inorganic phosphorus on the metabolic activity of bacteria in freshwater. Batch cultures with lake-water bacteria and dissolved organic carbon (DOC) extracted from a humic boreal river were arranged in an experimental matrix of three levels of exposure to simulated sunlight and three levels of phosphorus concentration. We measured an increase in bacterial biomass, a decrease in DOC and bacterial respiration as CO(2) production and O(2) consumption over 450 h. These measurements were used to calculate bacterial growth efficiency (BGE). Bacterial degradation of DOC increased with increasing exposure to simulated sunlight and availability of phosphorus and no detectable growth occurred on DOC that was not pre-exposed to simulated sunlight. The outcome of photochemical degradation of DOC changed with increasing availability of phosphorus, resulting in an increase in BGE from about 5% to 30%. Thus, the availability of phosphorus has major implications for the quantitative transfer of carbon in microbial food webs.     

Bacterioplankton in the littoral and pelagic zones of subtropical shallow lakes

We measured bacterioplankton (phylotypes detected by fluorescent in situ hybridisation, morphometric forms, abundance and production) in samples collected in summer in the littoral and pelagic zones of 10 subtropical shallow lakes of contrasting area (from 13 to 80,800 ha). Compared to the pelagic zones, the littoral zones were overall characterised by higher macrophyte dominance and lower concentrations of total phosphorus and alkalinity and higher concentrations of dissolved organic carbon (DOC) and humic substances. Similarities of bacterial production and biomass turnover and density of active phylotypes and morphotype proportions were related to similarities in a set of environmental variables (including nutrients, humic substances content, predator density and phytoplankton biomass), and some additionally to lake area. Horizontal heterogeneity in bacterioplankton variables (littoral versus pelagic) increased with lake area. Bacterioplankton biomass and production tended to be lower in the littoral zone than in the pelagic zone despite higher concentrations of DOC and humic substances. A likely explanation is higher predation on bacterioplankton in the littoral zone, although allelophatic effects exerted by macrophytes cannot be excluded. Our results indicate that organic cycling via bacterioplankton may be less efficient in the littoral zone than in the pelagic zone of shallow lakes.     

Effects of humic stress on the zooplankton from clear and DOC‐rich lakes

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How important is the crustacean plankton for the maintenance of water clarity in shallow lakes with abundant submerged vegetation?

• 1 We measured the abundance and biomass of filter‐feeding microcrustacean zooplankton and calculated their grazing impact on phytoplankton biomass during summer in five shallow, mesotrophic to eutrophic lakes. For three of the lakes data exist both from years with dense submerged vegetation and low turbidity (the clearwater state), as well as from years characterised by sparse vegetation and high turbidity (the turbid state). In the other two lakes data are available only for clearwater conditions.
• 2 In all lakes conditions of dense vegetation and clear water coincided with a low abundance of crustacean plankton during summer. In the three lakes that shifted, the calculated biovolume ingested by crustacean plankton (filtering rate) was 3–11 times lower during clearwater conditions compared with turbid conditions. Because phytoplankton biomass was lower during clearwater conditions, however, daily grazing pressure from microcrustacea (expressed as percentage of phytoplankton biomass) did not differ between states. In three of the five lakes, grazers were estimated to take less than 10% of the phytoplankton biomass per day, indicating filtration by zooplankton was not the most important mechanism to maintain clearwater conditions.
• 3 High densities of Cladocera were found in three of the lakes within dense stands of Charophyta. However, these samples were dominated by plant‐associated taxa that even during the night were rarely found outside the vegetation. This indicates that plant‐associated zooplankton has no major influence on the maintenance of water clarity outside the vegetation.
• 4 Spring peak abundance of Cladocera was observed in three of the lakes. In two of these, where seasonal development was studied in both the clearwater and the turbid state, spring peaks were lower during the clearwater state.
• 5 Predation, low food availability or a combination of both may explain the low zooplankton densities. Phytoplankton may be limited by low phosphorus availability in the lakes dominated by Charophyta. Our results indicate that the importance of zooplankton grazing may have minor importance for the maintenance of the clearwater state in lakes with dense, well‐established submerged vegetation.

     

Seasonal development of bacterioplankton in two forest lakes in central Sweden

Bacteria were counted with acridine orange epifluorescence technique in two humic lakes during 3 years. Less than 1% of the cells were found attached to detritus aggregates. 73% of the total number and 48% of the total volume were smaller than 1 µm. The mean cell volume ranged from 0.10 to 0.35 µm³ with the highest cell volumes occurring during early summer contemporarily with the growth of the bacterial biomass and probably indicating favourable growth conditions. The mean density of bacteria in oligotrophic brown-water lakes is higher than in oligotrophic clear-water lakes. The development of bacterial biomass showed a regular and seasonally dependent pattern with maxima during early summer and autumn. The importance of different factors for the regulation of bacterial biomass is discussed. Three different approaches were used to estimate bacterial production. These resulted in an average production rate of 15–60 µg C · l^?1 · d^?1 during the growing season. It was concluded that allochthonous sources comprised a significant part of the energy supply to the bacteria in the two humic lakes.     

Relationship between bacterial community composition and bottom-up versus top-down variables in four eutrophic shallow lakes

Bacterial community composition was monitored in four shallow eutrophic lakes during one year using denaturing gradient gel electrophoresis (DGGE) of PCR-amplified prokaryotic rDNA genes. Of the four lakes investigated, two were of the clearwater type and had dense stands of submerged macrophytes while two others were of the turbid type characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had high nutrient levels (total phosphorus, >100 micro g liter(-1)) while the other lakes had relatively low nutrient levels (total phosphorus, <100 micro g liter(-1)). For each lake, seasonal changes in the bacterial community were related to bottom-up (resources) and top-down (grazers) variables by using canonical correspondence analysis (CCA). Using an artificial model dataset to which potential sources of error associated with the use of relative band intensities in DGGE analysis were added, we found that preferential amplification of certain rDNA genes over others does not obscure the relationship between bacterial community composition and explanatory variables. Besides, using this artificial dataset as well as our own data, we found a better correlation between bacterial community composition and explanatory variables by using relative band intensities compared to using presence/absence data. While bacterial community composition was related to phytoplankton biomass in the high-nutrient lakes no such relation was found in the low-nutrient lakes, where the bacterial community is probably dependent on other organic matter sources. We used variation partitioning to evaluate top-down regulation of bacterial community composition after bottom-up regulation has been accounted for. Using this approach, we found no evidence for top-down regulation of bacterial community composition in the turbid lakes, while grazing by ciliates and daphnids (Daphnia and Ceriodaphnia) was significantly related to changes in the bacterial community in the clearwater lakes. Our results suggest that in eutrophic shallow lakes, seasonality of bacterial community structure is dependent on the dominant substrate source as well as on the food web structure.     

Short-term productivity responses of algae and bacteria to zooplankton grazing in two freshwater lakes

SUMMARY. 1. The specific productivities of algae and bacteria were measured in short-term (4 day) experiments consisting of enclosures with natural or reduced zooplankton biomass. Experiments were repeated five times over a season in each of two lakes that differed in the background concentration of dissolved organic carbon (DOC).
2. Algal biomass as estimated by chlorophyll a was suppressed in enclosures with ambient grazer levels in six of ten experiments and enhanced in one experiment. Distribution of chlorophyll among net and nanoplankton was not significantly affected by grazing.
3. Relative to enclosures with reduced zooplankton, normal grazer biomass (97–466μg 1⁻¹ dry weight) enhanced specific algal productivity in only one of five experiments in the low DOC take and had no effect in all five experiments in the high DOC lake. The main effects of grazers on algae was through removal of biomass rather than through indirect changes in turnover rate.
4. Between experiments, bacterial density was either unaffected, or mildly enhanced (4–87%) in enclosures with ambient macrozooplankton compared to those with reduced levels. Bacterial productivity and turnover estimated by incorporation of [³H]thymidine into DNA showed different responses across experiments; increasing, declining or remaining the same with grazer minipulation. This variability was not related to differences in dissolved primary production or to background DOC between lakes or experiments. Comparison of bacterial productivities based on thymidine incorporation rates with changes in cell densities indicated that control of bacterial loss processes by macrozooplankton is more important than control of growth rates.     

Dissolved organic carbon in a humic lake: effects on bacterial production and respiration

Allochthonous matter was the main source of carbon for pelagic bacteria in a humic lake, accounting for almost 90% of the carbon required to support observed bacterial growth. The estimated contribution from zooplankton excretion was of the same magnitude as direct phytoplankton release, both accounting for 5–7% of bacterial demands for dissolved carbon. Bacteria were an important source of carbon both for heterotrophic phytoplankton and for filter feeding zooplankton species, further stressing the role of humus DOC in overall lake productivity. The high contribution of allochthonous DOC implies a stoichiometry of dissolved nutrients with a surplus of C relative to P. The high P cell quota of bacteria suggest that under such conditions they are P-limited and act like net consumers of P. Excess C will be disposed of, and bacterial respiration rate will increase following a transition from carbon-limited bacterial growth towards mineral-nutrient-limited growth. Thus the high community respiration and frequent CO₂-supersaturation in humic lakes may be caused not only by the absolute supply of organic C, but also by the stoichiometry of the dissolved nutrient pool.     

Influence of humic substances on bacterial and viral dynamics in freshwaters

Bacterial and viral abundances were measured in 24 lakes with dissolved organic carbon (DOC) concentrations ranging from 3 to 19 mg of C liter(-1). In addition, a laboratory experiment was performed to test the effects of different sources of carbon (i.e., glucose and fulvic acids) and nutrients on the dynamics of viruses and bacteria. In the lake survey, no correlation was found between virus abundance and DOC concentration, yet there was a significant positive correlation between bacterial abundance and DOC concentration. A negative correlation was found between the virus-to-bacteria ratio and DOC level. These results are in agreement with our findings in the laboratory, where virus counts were significantly lower in treatments with fulvic acid additions than in a control (mean, 67.4% +/- 6.5% of the control). Virus counts did not differ significantly among the control and treatments with glucose, indicating that it was the type of organic carbon and not quantity which had an impact on viruses. Results from this study suggest that the way viruses control bacterial assemblages in humic lakes is different from the mechanism in clear water systems.