Winter limnology: a comparison of physical,chemical and biological characteristics in two temperate lakes during ice cover

The winter dynamics of several chemical, physical, and biological variables of a shallow, polymictic lake (Opinicon) are compared to those of a deep, nearby dimictic lake (Upper Rock) during ice cover (January to early April) in 1990 and 1991. Both lakes were weakly inversely thermally stratified. Dissolved oxygen concentration was at saturation (11–15 mg l⁻¹) in the top 3 m layer, but declined to near anoxic levels near the sediments. Dissolved oxygen concentrations in the deep lake were at saturation in most of the water column and approached anoxic levels near the sediments only. Nutrient concentrations in both lakes were fairly high, and similar in both lakes during ice cover. Total phosphorus concentrations generally ranged between 10–20 μg l⁻¹, NH₄-N between 16–100 μg l⁻¹, and DSi between 0.9–1.9 mg l⁻¹; these concentrations fell within summer ranges. NO₃-N concentrations were between 51–135 μg l⁻¹ during ice cover, but occurred at trace concentrations (<0.002 μg l⁻¹) during the summer. The winter phytoplankton community of both lakes was dominated by flagellates (cryptophytes, chrysophytes) and occasionally diatoms. Dinoflagellates, Cyanobacteria and green algae were poorly represented. Cryptophytes often occurred in fairly high proportions (20–80%) throughout the water column, whereas chrysophytes were more abundant just beneath the ice. Zooplankton population densities were extremely low during ice cover (compared to maximum densities measured in spring or summer) in both lakes, and were comprised largely of copepods.     

A method of collecting water samples from immediately below an ice cover

A plexiglas box with cellular neoprene gaskets in two opposite faces which can be frozen in situ in a developing ice cover on a lake is described along with a water sampling system using evacuated glass tubes (Vacutainers) and blood collecting needles. This system allows collecting of water samples from directly beneath the ice without disturbing the ice cover. The box may be removed in an ice block at the end of the sampling period.     

Winter behaviour of juvenile Atlantic salmon Salmo salar L. in experimental stream channels: effect of substratum size and full ice cover on spatial distribution and activity pattern

Activity and choice of areas offering different cover (substratum or surface ice) for juvenile Atlantic salmon Salmo salar were studied in experimental stream channels during winter. Channels were completely ice covered between December and March. During this period, the ice thickness increased from 50 to 300 mm after which 50% of the ice was experimentally removed and followed by c. 2·5-fold increase in discharge to simulate the effects of spring flood. Large substrata provided preferred habitats but areas with small substratum sizes were also used when full surface ice provided above-stream cover and the stream discharge was relatively low. The fish remained nocturnal throughout the study but the level of day activity significantly increased as the surface ice became thicker. Maximum movement distance during a 24 h period and homing-at-dawn behaviour remained at a constant level throughout the main winter, but significantly changed during the simulated spring flood (mean ± s . e . maximum extent of movements within 24 h increased from 1·1 ± 0·1 to 3·0 ± 0·5 m; homing behaviour decreased from the highest level of 89·3 to 34·6% during spring flood). Overwinter survival was high (92·9%). Relative mass increase during the study ranged from –8·3 to 28·5%, and 84% of the juvenile Atlantic salmon gained mass. The highest rates of mass increase were associated with frequent movements between areas of different substratum size. The results indicate that during winter: (1) Atlantic salmon parr preferred large substratum cover compared with surface ice cover at the fish densities studied here, (2) juvenile Atlantic salmon were predominantly nocturnal but diurnal activity increased as surface ice became thicker and (3) increase in water discharge during spring altered the behaviour of juvenile Atlantic salmon and may have caused additional habitat shifts.     

Microbial activity under the ice cover of the shallow Neusiedler See (Austria, Central Europe)

In an attempt to assess bacterioplankton production and growth yieldunder low temperature conditions and to compare bacterioplankton withphytoplankton production in the ice-covered water column of the shallowNeusiedler See, outdoor measurements under near in situ conditions wereperformed during the winter of 1995/96. During the investigation period,mean chlorophyll (Chl) a concentration was 21.03 ± 14.95 µg Chla l^-1. Phytoplankton primary production integrated over thewater column ranged from 1.35 to production integrated over the water columnranged from 1.35 to 4.23 mg C m^-2 d^-1 (mean± SD = 2.46 ± 1.06 mg C m^-2d^-1). Bacterial abundance varied from 20 to 40×10⁵ ml^-1 for most of the investigationperiod and increased by the end of March concomitantly with the increase intemperature from 1.3 to 6.3 °C within 5 days. Mean bacterial productionwas 15.3 ± 12.8 µg C l^-1 d^-1(range: 3.0 to 41.7 µg C l^-1 d^-1) and meanbacterial growth rate 0.23 ± 0.16 d^-1 following closelythe pattern in bacterial production. DOC concentration declined linearlyfrom 20.7 mg C l^-1 to 16.45 mg C l^-1 over the 4months period of ice cover. The contribution of humic substances to thetotal DOC pool declined from 43.6% at the end of November to37.3% at the end of March. Calculated on an area basis, phytoplanktonproduction amounted to only 16% of bacterial production which makesit unlikely that phytoplankton supply substrate for bacterioplankton growthin significant quantities when the lake is ice covered. From the observeddecline in DOC over the investigation period and assuming only negligibleinput of DOC from other sources we calculated an average DOC uptake by thebacterioplankton community of 47.5 µg C l^-1d^-1 resulting in a bacterial growth efficiency of 15.9%for the ice covered conditions. Based on the growth efficiency we estimatethat pelagic primary production amounts to 2.8% of the bacterialcarbon demand. This might indicate that the bacterioplankton in NeusiedlerSee sustain their high growth rates at low temperatures (<2°C formost of the investigation period) by using probably the DOC originating fromthe previous season. This DOM stems most likely from the decay of the reedPhragmites australis and its epiphytes and, probably of minor importance,from phytoplankton leachates.     

D1, but not D2, dopamine receptor regulates steroid levels during the final stages of pikeperch gametogenesis

In pikeperch, Sander lucioperca, aquaculture hormonal treatment is usually applied to synchronize ovulation. However, the effect of dopamine (DA) receptor antagonists, in particular those blocking the D1 DA receptors, remains unknown. Thus, the aim of the present study was to investigate and compare the effects of D1 and D2 DA receptor antagonists on the sex-steroid production and reproductive performance of the species. Two experiments were performed during which mature pikeperch females were injected with different molecules: NaCl 0.9% (negative control) or human chorionic gonadotropin 500 IU/kg (positive control) in both experiments, metoclopramide (a D2 receptor antagonist; 4 mg/kg or 20 mg/kg) or SCH23390 (a D1 receptor antagonist; 0.8 mg/kg or 4 mg/kg) alone (experiment 1) or in combination with a salmon gonadotropin-releasing hormone analogue (sGnRHa at 25 µg/kg; experiment 2). In experiment 2, fish were also injected with sGnRHa (25 µg/kg) as positive control. Samplings of oocytes and blood were performed on the day of injection and after 24 h (both experiments), after 48 h (experiment 2) and at the time of ovulation (both experiments). In non-ovulating fish, samplings were performed 7 days (experiment 1) or 14 days (experiment 2) after injection. In experiment 2, various zootechnical parameters of fertilized eggs were recorded (survival, hatching and malformation rates). The two antagonists alone were ineffective in inducing the final stages and regulating sex-steroid (testosterone, 11 ketotestosterone, 17β estradiol and 17,20β-dihydroxy-4-pregnen-3-one) production. When administered with sGnRHa, both SCH23390 and metoclopramide induced the final stages. However, only SCH23390 stimulated testosterone (4 mg/kg) and 17β estradiol (0.8 mg/kg) production compared with sGnRHa alone. None of the treatments affected the survival, hatching or malformation rates. This is the first report suggesting that in pikeperch the D1, but not the D2, DA receptor antagonist would be involved in the testosterone and 17β estradiol production as a potentiator of the sGnRHa effect.     

Seasonal sea ice cover as principal driver of spatial and temporal variation in depth extension and annual production of kelp in Greenland

We studied the depth distribution and production of kelp along the Greenland coast spanning Arctic to sub‐Arctic conditions from 78 ºN to 64 ºN. This covers a wide range of sea ice conditions and water temperatures, with those presently realized in the south likely to move northwards in a warmer future. Kelp forests occurred along the entire latitudinal range, and their depth extension and production increased southwards presumably in response to longer annual ice‐free periods and higher water temperature. The depth limit of 10% kelp cover was 9–14 m at the northernmost sites (77–78 ºN) with only 94–133 ice‐free days per year, but extended to depths of 21–33 m further south (73 ºN–64 ºN) where >160 days per year were ice‐free, and annual production of Saccharina longicruris and S. latissima, measured as the size of the annual blade, ranged up to sevenfold among sites. The duration of the open‐water period, which integrates light and temperature conditions on an annual basis, was the best predictor (relative to summer water temperature) of kelp production along the latitude gradient, explaining up to 92% of the variation in depth extension and 80% of the variation in kelp production. In a decadal time series from a high Arctic site (74 ºN), inter‐annual variation in sea ice cover also explained a major part (up to 47%) of the variation in kelp production. Both spatial and temporal data sets thereby support the prediction that northern kelps will play a larger role in the coastal marine ecosystem in a warmer future as the length of the open‐water period increases. As kelps increase carbon‐flow and habitat diversity, an expansion of kelp forests may exert cascading effects on the coastal Arctic ecosystem.     

Characterization of the gut microbiota in early life stages of pikeperch Sander lucioperca

This study characterized the gastrointestinal microbiome of nine juvenile farmed pikeperch Sander lucioperca using a metagenomics approach based on bacterial 16S rRNA gene sequencing. Potential changes in the gut microbiota during 2 months of S. lucioperca juvenile life were investigated. Results revealed that gut microbiota was dominated by Proteobacteria (95–92%), while other phyla Firmicutes (1–1·5%) and Actinobacteria (0·9–1·5%) were less abundant. At the family level, fish‐gut microbiota were dominated by Enterobacteriaceae, which constituted c. 83% of all DNA sequence reads. Such a situation was present in all of the examined fish except one, which showed a different proportion of particular microbial taxa than the other fish. In this fish, a higher relative abundance (%) of Fusobacteria (21·0%), Bacteroidetes (9·5%) and Firmicutes (7·5%) was observed. There were no significant differences in the gut microbiome structure at different stages of development in the examined fish. This may indicate that Proteobacteria inhabiting the gut microbiota at an early stage of life are a necessary component of the pikeperch microbiome that may support proper nutrition of the fish. The information obtained on the gut microbiome could be useful in determining juvenile S. lucioperca health and improving rearing conditions by welfare monitoring in aquaculture.     

Spectral transmittance of lake ice from 400–850 nm

Spectral transmittance signatures for a variety of freshwater ice types were collected in the 400–850 nm range. Clear ice showed the highest transmittances, snow-covered ice the lowest and other ice types intermediate transmittances. All of the ice spectral transmittance curves showed the same general shape with no mutually-exclusive characteristics exhibited by any ice type. The magnitude of the transmittance values was the primary distinguishing factor. The range of transmittance values directly under the ice was remarkably narrow (excluding snow-covered ice) and markedly different from values lower in the water column.     

Microbiota within the perennial ice cover of Lake Vida, Antarctica

Lake Vida, located in the McMurdo Dry Valleys, Antarctica, is an 'ice-sealed' lake with approximately 19 m of ice covering a highly saline water column (approximately 245 ppt). The lower portions of the ice cover and the lake beneath have been isolated from the atmosphere and land for circa 2800 years. Analysis of microbial assemblages within the perennial ice cover of the lake revealed a diverse array of bacteria and eukarya. Bacterial and eukaryal denaturing gradient gel electrophoresis phylotype profile similarities were low (<59%) between all of the depths compared (five depths spanning 11 m of the ice cover), with the greatest differences occurring between surface and deep ice. The majority of bacterial 16S rRNA gene sequences in the surface ice were related to Actinobacteria (42%) while Gammaproteobacteria (52%) dominated the deep ice community. Comparisons of assemblage composition suggest differences in ice habitability and organismal origin in the upper and lower portions of ice cover. Specifically, the upper ice cover microbiota likely reflect the modern day transport and colonization of biota from the terrestrial landscape, whereas assemblages in the deeper ice are more likely to be persistent remnant biota that originated from the ancient liquid water column of the lake that froze.     

Effects of snow cover on the benthic fauna in a glacier-fed stream

1. Alpine streams above the tree line are covered by snow for 6–9 months a year. However, winter dynamics in these streams are poorly known. The annual patterns of macroinvertebrate assemblages were studied in a glacial stream in the Austrian Alps, providing information on conditions under the snow.
2. Snow cover influenced water temperature, the content of benthic organic matter and insect development. Taxa richness and abundance of macroinvertebrates did not show a pronounced seasonal pattern. The duration of the autumn period with stable stream beds was important in determining the abundance and composition of the winter fauna.
3. There were significant differences in species composition between summer and winter. Two potential strategies in larval survival were evident: adaptation to the extreme abiotic conditions in summer (e.g. Diamesa spp.) or avoidance of these conditions and development during winter (e.g. Ephemeroptera and Plecoptera).
4. A comparison of a stream reach with continuous snow cover and a stream reach that remained open throughout winter showed that conditions under snow are suboptimal. At the open stream site, with higher water temperatures and greater food supply (benthic organic matter content), abundance and taxa richness was higher and larval growth was faster. Several taxa were found exclusively at this site.
5. Winter conditions did not provide an entirely homogeneous environment, abiotic conditions changed rapidly, especially at the onset of snowfall and at snowmelt. Continuous monitoring is necessary to recognize spatial and temporal heterogeneity in winter environments and the fauna of alpine streams.     

Genetic differentiation in pikeperch (Sander lucioperca): the relative importance of gene flow, drift and common history

To investigate the relative importance of homogenizing factors, such as gene flow, and diversifying factors, such as drift, genetic variation in pikeperch ( Sander lucioperca ) in two Fennoscandian regions (North and South) was analysed with microsatellites. Allelic richness and the degree of differentiation were significantly higher in the North ( F _ST= 0·20) than in the South ( F _ST= 0·064). In northern areas, assignments of genotypes were almost exclusively to the population of origin, but in southern areas, the proportion of correct assignments was significantly lower. Most samples exhibited significant heterozygote deficits, and the level of relatedness was higher than expected from randomness. These combined results suggest that there has been more gene flow between populations in southern areas than in northern areas, where the importance of genetic drift has been greater. Effective population sizes were small ( c. 100) and did not differ between areas. The effect of a common history appears minor, and thus processes such as genetic drift and gene flow have been more influential in shaping the patterns of genetic diversity in this species.     

Vertical and seasonal distributions of micro-organisms,zooplankton and phytoplankton in a eutrophic lake

The vertical distributions of bacteria, picoalgae,protozoan and metazoan zooplankton, and phytoplanktonin the highly eutrophic Lake Köyliönjärvi(SW Finland) were studied monthly during the period ofice-cover in January-April 1996. For comparison, wealso provide some data on the distributions of theplankton during the summer. The whole watercolumn remained oxic during the ice-covered period,although the near-bottom oxygen concentrations werealways very low. The heterotrophic nanoflagellateswere more abundant in winter than in summer, butciliates, picoalgae and bacteria were more numerous insummer. In general both zooplankton and phytoplanktonhad low biomass during the ice-covered period.However, the biomass of the diatom Aulacoseiraislandica ssp. islandica was high under the icein April. The calanoid copepod Eudiaptomusgraciloides was the dominant zooplankton species fromJanuary to March, but had almost disappeared by thebeginning of April and did not increase again until inJune. The dominant rotifer species in winterwere Keratella cochlearis, Filinia terminalis,and Filinia longiseta in the surface water andRotaria neptunia near the bottom.     

Effects of ice cover on the diel behaviour and ventilation rate of juvenile brown trout

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Photophysiology and albedo-changing potential of the ice algal community on the surface of the Greenland ice sheet

Darkening of parts of the Greenland ice sheet surface during the summer months leads to reduced albedo and increased melting. Here we show that heavily pigmented, actively photosynthesising microalgae and cyanobacteria are present on the bare ice. We demonstrate the widespread abundance of green algae in the Zygnematophyceae on the ice sheet surface in Southwest Greenland. Photophysiological measurements (variable chlorophyll fluorescence) indicate that the ice algae likely use screening mechanisms to downregulate photosynthesis when exposed to high intensities of visible and ultraviolet radiation, rather than non-photochemical quenching or cell movement. Using imaging microspectrophotometry, we demonstrate that intact cells and filaments absorb light with characteristic spectral profiles across ultraviolet and visible wavelengths, whereas inorganic dust particles typical for these areas display little absorption. Our results indicate that the phototrophic community growing directly on the bare ice, through their photophysiology, most likely have an important role in changing albedo, and subsequently may impact melt rates on the ice sheet.     

UV radiation and potential biological effects beneath the perennial ice cover of an antarctic lake

High-resolution spectral scans of solar ultraviolet radiation (UVR) were obtained directly beneath the 4.0–5.0 m thick, perennial ice cover of Lake Hoare, South Victoria Land, Antarctica. Both UVA (320–400 nm) and UVB (280–320 nm) radiation were detectable beneath the ice using a diver-deployed, underwater scanning spectroradiometer which permitted accurate measurement in the 280–340 nm range, while avoiding effects of surface shading and/or hole effects. UVR at wavelengths <310 nm was not detectable below the ice. This lower wavelength UVB appears to penetrate the Lake Hoare ice to depths of no more than 1.5 m during relatively cloud-free austral summer days. Based upon estimated biologically effective UVR dosages and DNA dosimeter data, exposure of benthic and planktonic microbes to the UVR encountered immediately beneath the ice is unlikely to inhibit microbial metabolism. Although waters of oligotrophic antarctic lakes are highly transparent to UVR, the thick, high scattering and optically dense ice covers on many of these lakes offers organisms a degree of protection largely unavailable in temperate and tropical systems. Thinning or complete loss of these overlying ice covers is likely to have major consequences for the structure of antarctic lake microbial communities.     

Studies of ice melting using molecular dynamics

In this paper, the melting of ice 1h is studied using molecular dynamics (MD). Common potential functions employed in the MD simulations include SPC/E, TIP4P, TIP5P, TIP4/ice and TIP5P/E. We first conducted melting of ice bulks and then studied the melting speed of the ice/water interface during ice melting. It is found that various potentials result in similar ice-melting phenomena. The result is compared with the analytical solution for phase change problem. We also studied size effects and temperature gradient effects on ice melting.     

A device for remote sampling of benthic algae under ice

A benthic algae sampling device designed for use through surface ice, uses the principle of pivoting arms. It is designed to be activated below ice sheets and is capable of passing easily through a hole of 11 cm diameter bored by a SIPRE ice auger.     

Effects of hanging ice dams on winter movements and swimming activity of fish

Formation of a hanging dam, a large, thick sub-surface accumulation of frazil ice that blocked river flow, altered physical environmental conditions dramatically and common carp Cyprinus carpio , and brown trout Salmo trutta , evacuated the pool in which the dam developed. A mean of 80·1% of the pool volume was filled with frazil ice. Mean and bottom water velocities in the pool increased from 6 to 27 cm s⁻¹ and from 4 to 21 cm s⁻¹, respectively, when the hanging dam formed, and water depth decreased from a mean of 2·25 to 0·45 m. Activity levels of fish leaving the pool varied. Two carp had higher swimming activities when in the over wintering pool, compared to the period when they were out of it, and a third showed opposite results.     

Winter movements and habitat use of riverine brown trout, white sucker and common carp in relation to flooding and ice break-up

Movement of radio-tagged riverine brown trout Salmo trutta , white sucker Catostomus commersoni and common carp Cyprinus carpio in the Grand River, Ontario, was minimal during the winter except during periods of high water discharge or the break-up of river ice. Ice break-up and flooding occur when southern air masses penetrate northwards in winter and as air temperatures increase in spring. Both white suckers and common carp moved the longest distances, generally in a downstream direction, during flooding and ice break-up. Brown trout moved less during these times. All three species had positive relationships between distance moved and water discharge. During periods of high water discharge and ice break-up, white suckers and common carp used backwater habitats more and main channel habitats less than during periods of low water discharge. During high flows, brown trout tended to use runs more and pools less. The break-up of river ice and accompanying flooding resulted in several fish being stranded on the floodplain. Backwater habitats appear to be important areas of refuge for riverine fishes during periods of flooding and ice break-up.     

The use of cover crops in cereal-based cropping systems to control nitrate leaching in SE England

Field experiments were done to evaluate the extent to which cover crops can be used to help farmers comply with current legislation on nitrate leaching from arable land in nitrate vulnerable zones. Nitrate leaching was measured in sandy loam and chalky loam soils under a range of early sown (mid-August) cover crops at two sites in SE England, and in the subsequent winter following their incorporation. Cover crop species tested were forage rape, rye, white mustard, a rye/white mustard mixture, Phacelia and ryegrass. Additional treatments were weeds plus cereal volunteers, a bare fallow and a conventional winter barley crop sown one month later than the cover crops and grown to maturity. Cover crop and bare fallow treatments were followed by spring barley. This was followed by winter barley, as was the conventional winter barley crop. In the winter immediately after establishment, early sown cover crops decreased nitrate leaching by 29–91% compared to bare fallow. They were most effective in a wet winter on the sandy loam where nitrate leaching under bare fallow was greatest. There was little difference between cover crop species with respect to their capacity to decrease nitrate leaching, but losses were consistently smaller under forage rape. The growth of weeds plus cereal volunteers significantly decreased nitrate leaching on the sandy loam compared with a bare fallow, but was less effective on the chalky loam. Nitrate leaching under the later sown winter barley was often greater than under cover crops, but under dry conditions leaching losses were similar. In the longer-term, in most cases, the inclusion of cover crops in predominantly cereal-based cropping systems did not significantly decrease cumulative nitrate leaching compared with two successive winter cereals. In summary, early sown cover crops are most likely to be effective when grown on freely drained sandy soils where the risk of nitrate leaching is greatest. They are less likely to be effective on poorer drained, medium-heavy textured soils in the driest parts of SE England. In these areas the regeneration of weeds and cereal volunteers together with some additional broadcast seed may be sufficient to avoid excessive nitrate losses. In the short-term, mineralization of N derived from the relatively small cover crops grown once every 3–4years in cereal-based cropping systems is unlikely to contribute greatly to nitrate leaching in later years and adjustments to fertilizer N recommendations will not usually be necessary.