Nutrient limitation of bacterioplankton and phytoplankton in humic lakes in northern Sweden

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L^–1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), respectively. A third lake was unfertilized and served as a control. In addition to this lake fertilization experiment, data from different regional surveys were used to assess the role of different limiting factors.
2. The P fertilization had no effects on bacterioplankton or phytoplankton, while phytoplankton were significantly stimulated by N fertilization. Inorganic nutrient limitation of bacterioplankton was a function of DOC concentration in water of the investigated region and nutrient-limited bacteria were found only in lakes with DOC concentrations less than around 15 mg L^–1
3. The fertilization experiments demonstrated that the DOC-rich experimental lakes contained a bioavailable pool of P that was not utilized to its full potential under natural conditions. The overall mobilization of energy (bacterioplankton plus phytoplankton) in the experimental lakes was restricted by lack of inorganic N.     

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.     

Nutrient budgets and the regulation of nutrient concentrations in a small sub-arctic lake in northern Sweden

SUMMARY. A 5-year study was made of the hydrology and water chemistry in the watershed of Lake Stugsjön, a small lake in the sub-arctic region of northern Sweden. Of the total yearly runoff, 80% occurs during the spring thaw in May and June and largely regulates the nutrient loading. The ice-free period of the lake is June—October. Atmospheric deposition of nitrogen and phosphorus was among the lowest recorded anywhere, in spite of which it accounts for approximately half of the total summer nutrient loading. Dry fall-out made up the major part of the total atmospheric deposition of nitrogen and phosphorus during the summer.
Conductivity measurements showed that the major constituents were not influenced by processes within the lake.
The nitrogen concentration in the lake during the summer was probably regulated by benthic algae which assimilated inorganic nitrogen from the sediments and released organic nitrogen to the lake water.
Phosphorus loading and water temperature in combination showed a clear relationship with the mean concentration of chlorophyll-α during the ice-free period.     

Vertical distributions of bacteria and algae in a steeply stratified humic lake under high grazing pressure from Daphnia longispina

The vertical distributions of bacteria and algae in a steeply stratified, highly humic lake were studied during three 24 h periods in summer. The highest bacterial and algal densities and biomasses were recorded in the anoxic hypolimnion. The bacterial biomass in the hypolimnion was composed mainly of photosynthetic green sulphur bacteria (Chlorobium) which occurred at very low light intensity (< 1.5 μmol m⁻² s⁻¹). The numbers and biomasses of bacteria, both in the epilimnion and hypolimnion, were mostly higher at night than during the day, indicating possible asynchrony between the production and loss of bacteria. Because of vertical migration, the diurnal vertical distribution of algae was more variable than that of bacteria. Particularly in July and August, when cryptomonads were abundant, the biomass of algae was much higher in the epilimnion during the day than at night. The flagellated chlorophytes, Chlamydomonas spp. and Scourfieldia cordiformis, stayed mainly in the upper hypolimnion close to the oxic-anoxic boundary zone where only a small proportion of Daphnia longispina was continuously present. Unpalatable Mallomonas chrysophytes with silicified plates and bristles, and small, presumably heterotrophic, flagellates stayed in the oxic epilimnion together with a dense (up to 300 ind l⁻¹) population of D. longispina. The results indicated that, besides the physical and chemical properties of the water column, grazing pressure by Daphnia longispina strongly affected the vertical distribution of microorganisms in this polyhumic lake.     

Diel patterns of zooplankton grazing in a shallow lake

A diel survey of in situ species-specific zooplankton clearancerates (with radioactively labelled Chlorella cells) was conductedin the shallow lake of Cr?teil characterized by small-sizedplanktonic forms (algae<10µm and zooplankton <1.3mm). Experiments were performed every 4h at two depths (1 and4m). Power functions relating individual filtering rates tobody length were established for the three most abundant cladoceransand for calanoids (nauplii being included in this feeding group),for each depth and time. No filtering penodicity was observedin Ceriodaphnia spp., adults and copepodites of Eudiaptomusgracilis and Eurytemora velox, and nauplii. On the contrary,clear nocturnal filtering peaks were obtained for Daphnia spp.and for Diaphanosoma brachyurum, these being more pronouncedfor the larger individuals at 1 m deep. The observed diel periodicitycannot be explained by variations in physico-chemical parametersor food concentration alone. The ecological significance ofthis phenomenon in the polymictic lake of Cr?teil is discussedin the light of previously published data and the hypothesesaccounting for it.     

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.     

Planktonic food chains of a highly humic lake

The development and metabolism of epilimnetic plankton from a highly humic lake was followed in late summer, when the predominant zooplankton species, Daphnia longispina, was very abundant (ca. 200 ind. l^?1). The experiment was made in two tanks: one with an unaltered plankton assemblage and one with larger zooplankton removed. The scarce phytoplankton community was also simple, consisting mainly of one Cryptomonas and two Mallomonas species. The abundance and species composition of smaller plankton was heavily influenced by grazing of Daphnia. In particular, the biomass, of heterotrophic flagellates increased after the removal of Daphnia. The biomass and production of bacterioplankton were not affected, and remained several times higher than that of phytoplankton. Bacterial production and grazing on bacteria were balanced, and when Daphnia was removed its grazing activity was compensated by flagellates. The removal of Daphnia did not affect the respiration or community net production of plankton. Among organisms smaller than zooplankton, bacteria seemed to be responsible for most of the respiration. The community net production was consistently negative even at the water surface, indicating an allochthonous carbon source. The results suggest that phytoplankton primary production was insufficient for the secondary production in the epilimnetic water of the study lake. The food requirements of bacteria and zooplankton, as well as of flagellates, each exceeded that supplied by phytoplankton primary production. The simple food chains in this experiment made it possible to reveal the functioning of the community so completely that dissolved organic matter is certainly comparable to or exceeds the importance of phytoplankton primary production as an energy and carbon source for food webs in this humic lake.     

Productivity of Daphnia longispina in a highly humic boreal lake

The development of the Daphnia longispina (O. F. Müller)population in a highly humic boreal lake was followed throughoutone growing season, and the amount of secondary production wasestimated in relation to primary production and available foodresources. The growth rate method was applied in the secondaryproduction measurements. Daphnia longispina did not appear inthe water column until 16 May, after which the animals werepresent throughout the growing season. The population showedthree density peaks; the first appeared in early June, and thesecond and third in mid-July and at the beginning of September,respectively. Somatic production followed a seasonal pattern,with highest production rates in midsummer. The maximum valueof 127 mg C m^–2day^–1 was recorded at the beginningof July. The total annual net production of D. longispina was7.9 g C m^–2. During most of the growing season, the primaryproductivity in the lake was well below 100 mg C m^–2 day^–1and the total annual productivity of photosynthetic algae was5.0 g C m^–2. We conclude that in this lake the zooplanktonpopulation did not rely on phytoplankton primary productionas a sole carbon source, but that most of the carbon must haveoriginated from bacterial production either directly or througha microbial loop.     

Zooplankton grazing on bacteria and cyanobacteria in a eutrophic lake

During in situ experiments in a cyanobacteria- and copepod-dominatedeutrophic subtropical lake, all taxa of protozoa and metazoanzooplankton grazed fluorescently labelled bacteria and all metazoanzooplankton taxa grazed large filamentous and colonial cyanobacteria.     

Seasonal dynamics,typhoons and the regulation of lake metabolism in a subtropical humic lake

1. We used high‐frequency in situ dissolved oxygen measurements to investigate the seasonal variability and factors regulating metabolism in a subtropical alpine lake in Taiwan between May 2004 and October 2005, specifically exploring how the typhoon season (from June or July to October) affects lake metabolism. 2. Gross primary production (GPP) and ecosystem respiration (R) both peaked in early summer and mid‐autumn but dropped during the typhoon season and winter. Yuan‐Yang Lake is a net heterotrophic ecosystem (annual mean net ecosystem production ?39.6 μmole O₂ m^?3). 3. Compared to the summer peaks, seasonal averages of GPP and R decreased by approximately 50% and 25%, respectively, during the typhoon season. Ecosystem respiration was more resistant to external disturbances than GPP and showed strong daily variation during typhoon seasons. 4. Changes in the quality and quantity of dissolved organic carbon controlled the temporal dynamics and metabolic regulation. External disturbances (typhoons) caused increased allochthony, increasing DOC and water colour and influencing lake metabolism. 5. Seasonal winter mixing and typhoon‐induced water mixing in summer and autumn play a key role in determining the extent to which the lake is a seasonal carbon sink or source to the atmosphere.     

Bacterial population dynamics in a meromictic lake.

Polyclonal antibodies against nine different bacteria isolated from Lake Saelenvannet in western Norway were produced, and the population dynamics of these strains in the lake were monitored through two spring seasons by immunofluorescence staining. The total counts of bacteria varied over time and space from 1.5 x 10(6) to 1.5 x 10(7) cells ml-1. The counts of specific bacteria were in the range of 10(3) to 10(4) cells ml-1 or less; in sum, they generally made up less than 1% of the bacterial community. Some populations showed significant changes in abundance, with blooms lasting 1 to 3 weeks. The rate of change (increase and decrease) in abundance during blooms was estimated to be 0.2 to 0.6 day-1. The average virus-to-bacteria ratio was 50, and there was a significant correlation between the abundances of virus and bacteria. Both protozoan grazing and lytic virus infection were assessed as possible mechanisms driving the variations in bacterial population density.     

Bat species diversity in a lake archipelago in central Sweden

The bat fauna of 35 islands in a large lake in central Sweden were examined using ultrasound detectors. We tested the hypothesis that there is no difference in species number between the mainland and the island fauna. Eight species were found. Species numbers were analysed against island area, area of some habitats (coniferous forest, deciduous forest, semi-open habitats and open habitats), degree of isolation (distance from mainland and from stepping stones) and time spent searching for bats. Species number increased with area of deciduous forest. Presence of houses tended to increase species number. There seems to be a negative relationship between species number and degree of isolation (nearly significant). The results suggest that at least three species, Myotis brandti (Eversmann, 1845), M. mystacinus (Kuhl, 1819) and Plecotus auritus (Linnaeus, 1758), are negatively affected by forest patchiness. These species occurred mainly on large islands. Thus, the results do not support the hypothesis. The reasons why some species avoid open habitats are discussed.     

Waterfowl grazing effects on submerged macrophytes in a shallow mediterranean lake

Waterfowl exclusion cages were set up in Sentiz Lake, an eutrophic shallow lake in León (NW of Spain) in order to determine the role of waterfowl herbivory on macrophyte biomass and species composition. Total macrophyte biomass was high during the study (250 g DW m⁻² in summer). The macrophyte community was mainly formed by Myriophyllum alterniflorum (95% cover), Ceratophyllum demersum (5%) and Potamogeton gramineus (<0.5%). High densities of co-occurring coots (Fulica atra; 24 ind/ha) and ducks (Anas penelope, A. strepera and A. platyrhynchos; 18 ind/ha) did not have a significant effect on macrophyte biomass in the lake. There were no statistical differences between total biomass inside and outside the exclosures, although plant biomass reached a higher value inside the cages than in the lake. Biomass species composition was significantly different inside and outside exclosures; C. demersum was more abundant in the cages than in the lake. P. gramineus, almost absent in the lake, became co-dominant with M. alterniflorum in some exclosures. The detailed study of M. alterniflorum flower buds in summer showed significant herbivory by coots. Flower bud abundance was lower in the lake (35% lower in June; 85% lower in July) than under waterfowl exclusion. The effect of waterfowl on macrophyte biomass in Mediterranean wetlands seems to be negligible as compared to effects identified in northern European lakes. Apart from an important role in dispersal, waterfowl in Mediterranean areas have a strong qualitative effect on the structure of plant communities by selecting most palatable species or their reproductive structures.     

Contribution of grazing in phytoplankton overall losses in a shallow French lake

SUMMARY 1. Based on data for ¹⁴C-primary production and biomass changes in a small and shallow lake (Créteil Lake, France), overall phytoplankton losses were calculated through an annual cycle (November 1985-October 1986). The summer period in 1986 is compared with two other summer periods in 1985 and 1980, these two years corresponding to extreme levels of algal biomass.
2. Independent from the trophic state of the lake, phytoplankton populations were dominated by small-sized species (<20 μm); their high growth rate (maximal in May and June: 0–8 day⁻¹) was characteristic of nanoplanktonic natural populations.
3. The positive correlation between phytoplankton losses and production indicates a close coupling between growth and loss processes.
4. With a high filtering rate (0.22 day⁻¹ as an annual average), zoo-plankton impact is considerable at any time of the year but especially in late summer, when grazing losses exceeded primary production.
5. Despite the uncertainty concerning the meaning of ¹⁴C-primary production, the persistence of small algae throughout the year implies that a great part of the phytoplankton production was harvested by grazers which led to a recycling of organic matter within the water column.     

Vertical and temporal heterogeneity of planktonic ciliated protozoa in a humic lake

Seasonal population dynamics and the vertical distribution ofciliates were studied in relation to the particular food resourcesoccurring in a humic and moderately acidic lake (Lake Vassivière).The abundance (1.4 x 10³–20.4 x 10³ cells l^–1 mean= 4.8 x 10³ cells l^–1) and biomass (0.5–34.6 µgC l^–1, mean = 6.0 µg C l^–1) of ciliated protozoawere low and close to values reported for oligotrophic environments.The species composition of the population varied greatly withdepth. Whereas large-sized species of oligotrichs, some of whichwere mixotrophic, dominated at the surface, haptorids were bestrepresented in deep waters. The spatial distribution of thevarious groups of ciliates was largely determined by light andthe vertical distribution of microbial food resources (detritus,bacteria, algae) within the water column of this brown-coloredlake.     

Factors determining the nutritive status and production of zooplankton in a humic lake

Zooplankton egg production and nutritional status as relatedto environmental factors were studied in a humic lake. Bothparameters exhibited a pronounced spring peak, a mid-summerdepression and a second, smaller autumnal peak. The overallmacrozooplankton community apparently suffered from severe foodlimitation, particularly with regard to food quality, duringthe study period. This limitation was confirmed with length/carbonregressions of well-fed and starved laboratory individuals ofDaphnia. Body carbon in the lake animals corresponded to thatof laboratory-raised, starved individuals. Regression analysisindicated that bacteria, particulate organic carbon and particulatenitrogen best explained changes in egg production, while algalbiomass was of negligible importance for all species. For Daphnia,size-specific carbon content (length/carbon regressions) andC/N ratios were computed for all sampling dates. a and b fromthe equation W=aL^b were positively correlated (P=0.001). a rangedover 1.17–4.44 and was positively correlated with size-specificcarbon content (P<0.001) and the C/N ratio (P=0.01). Whena mean time lag of 12 days was introduced, a was positivelycorrelated with egg production (P=0.04.6). b ranged over 2.05–3.54and was inversely correlated to the same parameters. Both simpleand multiple regression analysis confirmed that bacteria, particulateorganic carbon and particulate nitrogen gave the major positivecontribution to variance in size-specific C content, a and C/Nratio in Daphnia, whereas total zooplankton biomass was themain negative contributor to the same parameters.     

Feeding of Boeckella gracilipes (Copepoda, Calanoida) on ciliates and phytoflagellates in an ultraoligotrophic Andean lake

The calanoid copepod, Boeckella gracilipes, is the dominantcrustacean zooplankton in South Andean deep ultra-oligotrophiclakes. Combining field and experimental data we explored thefeeding of the copepod and its access to the mixotrophic ciliate,Ophrydium naumanni, in Lake Moreno Oeste (Patagonia, Argentina).Phytoplankton was dominated by nanoflagellates throughout thewater column. Ophrydium naumanni, which accumulates much ofthe chlorophyll a, as do copepodites and adults of B. gracilipes,has a deep distribution during the day, with maximal abundancesaround 30 m depth. Mouth-part morphology analysis of B. gracilipesindicated that the copepod has an omnivorous diet. Laboratoryexperiments showed that B. gracilipes could access O. naumannionly when it is offered as a single food item. However, whennatural phytoplankton and ciliate assemblages (including O.naumanni)are offered, B. gracilipes did not eat Ophrydium and preyedon the oligotrich, Strombidium viride, and phytoflagellateslike Chrysochromulina parva. The range of ingested sizes wasbroad (3.9–33 µm of equivalent spherical diameter)but all selected particles were motile ones with distinctivemovements, which would enhance the copepod particle detection.     

Early post-fire tree regeneration in a Picea-Vaccinium forest in northern Sweden

Abstract. Age and size structure of saplings of Picea abies, Pinus sylvestris and Betula pubescens were examined in a 26-yr old forest fire area in a Picea abies-Vaccinium myrtillus forest in northern Sweden. Picea, which is a shade-tolerant species, had its maximum regeneration prior to the shade-intolerant Pinus. The shift from Picea to Pinus regeneration in the late 1970s, did not seem to be related to variations in summer temperature. Instead, it is suggested that Picea established in the shade created by dead trees, and, that increased reindeer browsing of Betula in combination with a simultaneous thinning of the tree layer, favoured Pinus recruitment. These regeneration patterns do not confirm conventional views of post-fire succession in Sweden.     

Intrinsic breaking of internal solitary waves in a deep lake

Based on simulations with the Dubreil-Jacotin-Long (DJL) equation, the limiting amplitude and the breaking mechanisms of internal solitary waves of depression (ISWs) are predicted for different background stratifications. These theoretical predictions are compared to the amplitude and the stability of the leading internal solitary waves of more than 200 trains of ISWs observed in the centre of a sub-basin of Lake Constance. The comparison of the model results with the field observations indicates that the simulated limiting amplitude of the ISWs provides an excellent prediction of the critical wave height above which ISWs break in the field. Shear instabilities and convective instabilities are each responsible for about half of the predicted wave breaking events. The data suggest the presence of core-like structures within the convectively unstable waves, but fully developed and stable cores were not observed. The lack of stable trapped cores in the field can be explained by the results from dynamic simulations of ISWs with trapped cores which demonstrate that even slight disturbances of the background stratification cause trapped cores to become unstable.     

Composition and variations in the occurrence of dissolved free simple organic compounds of an unproductive lake ecosystem in northern Sweden

Low molecular weight organic carbon compounds are potentially important carbon and energy substrates to heterotrophic production in the aquatic environment. We studied the occurrence of dissolved free amino acids (AA), monosaccharides (CHO), and carboxylic acids (CA) in the subarctic Lake Diktar-Erik. The lake is unproductive with slightly humic water, and receives water via one major inlet stream draining a birch forest environment. The concentration of dissolved organic carbon (DOC) in the inlet stream was strongly correlated with the discharge. This relationship changed from season to season, indicating changes in the sources of the DOC entering the stream. AA and CHO each accounted for an average of less than 0.5% of the DOC. After high discharge events during the ice-free period, AA and CHO occurred in especially high concentrations. CA occurred in higher concentrations during the ice-free period, when it generally accounted for 20–30% of the DOC pool. The CA content relative to the total DOC pool was strongly inversely correlated with overall DOC concentration, and at low DOC levels the relative content of CA was high and vice versa. This followed a seasonal trend, with CA accounting for a smaller proportion of the DOC in winter and a larger part in spring/early summer. A conservative estimate suggested that the studied simple organic carbon compounds potentially could cover 30% of the bacterial gross production in the lake and therefore potentially also was an important source of CO₂ that occur in supersaturated concentrations in the lake.