Ace Lake: three decades of research on a meromictic,Antarctic lake

Ace Lake (Vestfold Hills, Antarctica) has been investigated since the 1970s. Its close proximity to Davis Station has allowed year-long, as well as summer only, investigations. Ace Lake is a saline meromictic (permanently stratified) lake with strong physical and chemical gradients. The lake is one of the most studied lakes in continental Antarctica. Here, we review the current knowledge of the history, the physical and chemical environment, community structure and functional dynamics of the mixolimnion, littoral benthic algal mats, the lower anoxic monimolimnion and the sediment within the monimolimnion. In common with other continental meromictic Antarctic lakes, Ace Lake possesses a truncated food web dominated by prokaryote and eukaryote microorganisms in the upper aerobic mixolimnion and an anaerobic prokaryote community in the monimolimnion, where methanogenic Archaea, sulphate-reducing and sulphur-oxidizing bacteria occur. These communities are functional in winter at subzero temperatures, when mixotrophy plays an important role in survival in dominant photosynthetic eukaryotic microorganisms in the mixolimnion. The productivity of Ace Lake is comparable to other saline lakes in the Vestfold Hills, but higher than that seen in the more southerly McMurdo Dry Valley lakes. Finally, we identify gaps in the current knowledge and avenues that demand further investigation, including comparisons with analogous lakes in the north polar region.     

Virus and microbial loop dynamics over an annual cycle in three contrasting Antarctic lakes

1. Viral and microbial loop dynamics were investigated over an annual cycle in three contrasting saline Antarctic lakes – Highway Lake (salinity 4‰), Pendant Lake (salinity 19‰) and Ace Lake, a meromictic system (with a mixolimnion salinity of 18‰) in order to assess the importance of viruses in extreme, microbially dominated systems. 2. Virus like particles (VLP) showed no clear seasonal pattern, with high concentrations occurring in both winter and summer (range 0.89 × 10⁷ ± 0.038 to 12.017 × 10⁷ ± 1.28 mL^?1). VLP abundances reflected lake productivity based on chlorophyll a concentrations. Bacterial abundances and biomass did not correlate with VLP numbers except in Pendant Lake, the most productive of the three lakes studied. 3. Pendant Lake supported the highest bacterial biomass (range Highway: 18.44 ± 1.35 to 59.43 ± 2.80 ng C mL^?1; Ace: 14.42 ± 2.69 to 68.39 ± 2.95 ng C mL^?1; Pendant: 31.36 ± 3.94 to 115.95 ± 4.49 ng C mL^?1) so that virus to bacteria ratios (VBR) (range 30.48 ± 7.96 to 96.67 ± 8.21) were higher in Ace Lake (range 30.58 ± 3.98 to 80.037 ± 1.60) and Highway Lake (range 18.63 ± 3.12 to 126.74 ± 6.50). 4. Negative correlations occurred between VLP and cryptophytes (dominant phototrophic nanoflagellates), suggesting that they were not hosts to lytic viruses. Among the other protists only the heterotrophic nanoflagellates of Highway Lake (dominated by the marine choanoflagellate Diaphanoeca grandis) showed a positive correlation with VLP. 5. The VLP was negatively correlated with photosynthetically active radiation (PAR) and temperature, both of which increased with ice thinning and breakout, increasing viral decay. In winter VLP probably persisted in cold, dark water. 6. High VLP concentrations and high VBR (values at the upper end of those reported for marine and lacustrine systems) indicated that viruses, most of which were probably bacteriophage, are a major element within the microbial communities in extreme, saline lakes.     

Flagellate nutritional versatility as a key to survival in two contrasting Antarctic saline lakes

1. Seasonal patterns of grazing and photosynthesis were investigated in two saline Antarctic lakes (Highway and Ace) in the Vestfold Hills (68°S). The phototrophic nanoflagellate (PNAN) community was dominated by Pyramimonas gelidicola and two morphological forms of a cryptophyte species that occurred throughout the year. Both species were mixotrophic on bacteria, and in Highway Lake they also exploited dissolved organic carbon as determined by the uptake of fluorescently labelled dextrans. 2. Clearance rates ranged between 0.02 and 0.21 nL h^?1 cell^?1 in Ace Lake and 0.004–1.05 nL h^?1 cell^?1 in Highway Lake. On occasion cryptophyte grazing equalled that of the heterotrophic nanoflagellates (HNAN). 3. Photosynthetic rates showed similar trends in both lakes, but there were differences in chlorophyll a specific rates and photosynthetic efficiency, probably related to the meromictic characteristic of Ace Lake. Primary production was measurable in winter and peaked in summer following the maxima of mixotroph grazing. 4. The HNAN community of Highway Lake achieved clearance rates of 0.02–1.80 nL h^?1 cell^?1 and removing between 50 and 693 ng bacterial carbon L^?1 day^?1, with highest impact in winter when HNAN were most abundant. The HNAN also ingested fluorescently labelled dextrans showing a preference for 4 and 500 kDa molecules. The more diverse HNAN community of Ace Lake had lower clearance rates (0.04–0.37 nL h^?1 cell^?1) and exerted a lower grazing pressure on bacterioplankton. In Highway Lake, where the HNAN community was dominated by the choanoflagellate Diaphanoeca grandis, there was a significant correlation between mean cell volume and clearance rate. 5. The major feature was that the microbial plankton functioned throughout the year by employing nutritional versatility.     

A comparison of phototrophic bacteria in two adjacent saline meromictic lakes

Two adjacent saline, meromictic lakes in Saskatchewan host different populations of phototrophic bacteria. Deadmoose Lake hosts a population of Lamprocystis roseopersicina (Chromatiaceae) while in Waldsea Lake a population of a Chlorobium species (Chlorobiaceae) is dominant. Differences in light quantity, light quality, temperature, pH and Lamprocystis' capacity for photoorganoheterotrophic growth explain why different genera of phototrophic bacteria are found within the two lakes. These phototrophic bacteria make a significant contribution to total photosynthetic productivity, fixing 14.3 and 32 g C m^-2 year ^-1 in Deadmoose and Waldsea Lake respectively.     

Viral dynamics and patterns of lysogeny in saline Antarctic lakes

Antarctic lakes are extreme ecosystems with microbially dominated food webs, in which viruses may be important in controlling community dynamics. A year long investigation of two Antarctic saline lakes (Ace and Pendant Lakes) revealed high concentrations of virus like particles (VLP) (0.20–1.26 × 10⁸ ml⁻¹), high VLP: bacteria ratios (maximum 70.6) and a seasonal pattern of lysogeny differing from that seen at lower latitudes. Highest rates of lysogeny (up to 32% in Pendant Lake and 71% in Ace Lake) occurred in winter and spring, with low or no lysogeny in summer. Rates of virus production (range 0.176–0.823 × 10⁶ viruses ml⁻¹ h⁻¹) were comparable to lower latitude freshwater lakes. In Ace Lake VLP did not correlate with bacterial cell concentration or bacterial production but correlated positively with primary production, while in Pendant Lake VLP abundance correlated positively with both bacterial cell numbers and bacterial production but not with primary production. In terms of virus and bacterial dynamics the two saline Antarctic lakes studied appear distinct from other aquatic ecosystems investigated so far, in having very high viral to bacterial ratios (VBR) and a very high occurrence of lysogeny in winter.     

A Limnological Survey of Lake Mainit,Philippines

Morphology, bathymetry and climate are described for Lake Mainit, one of the principal lakes of Southeast Asia. Thermal and water chemistry data show that the lake is not meromictic, despite its great depth, and that the water is dilute. Primary production is high but standing crop and diversity of phytoplankton and zooplankton range from moderate to low. The species composition of plankton and other biota is discussed.     

Life and Times of Five Saskatchewan Saline Meromictic Lakes

Five closed saline lakes near Humboldt, Saskatchewan, were found to be meromictic. Two of these lakes (Waldsea, Deadmoose) were first discovered to be meromictic in the early 1970s and three (Arthur, Marie, Sayer) in 1985. The origin of their meromixis is ectogenic. One of the lakes, Waldsea, had surface salinities far higher in 1960–1961 than those of 1970 or later and as high as that of the monimolimnion occasionally was from 1970 to the present. During the late 1960s to 1980 the lake level of Waldsea rose four metres as a result of higher than normal snowpacks and subsequent high snowmelt runoff. Endogenic processes of freezing out of salts from the upper metre during ice formation and precipitation of sodium sulphate during autumn cooling also promote meromixis. The lakes which are located in depressions in a relatively flat topography are very exposed to periodic high velocity westerly winds. Although Deadmoose and Waldsea lakes are relatively deep, Arthur, Marie and Sayer lakes have maximum depths of only three to five metres. Meromixis has persisted until the present in three lakes but Marie and Arthur lakes became holomictic during the autumn of 1988, a severe drought year. Bacterial plates were prominent in Waldsea, Deadmoose and Sayer lakes. BChl-a and BChl-d were present in 1988 with maxima of 2652 mg · m⁻³ BChl-a and 4290 mg · m⁻³ BChl-d in Sayer Lake. BChl-a virtually disappeared in subsequent years.     

Viruses in the plankton of freshwater and saline Antarctic lakes

1. Virus‐like particle (VLP) abundances in nine freshwater to saline lakes in the Vestfold Hills, Eastern Antarctica (68° S) were determined in December 1999. In the ultra‐oligotrophic to oligotrophic freshwater lakes, VLP abundances ranged from 1.01 to 3.28 × 10⁶ mL^–1 in the top 6 m of the water column. In the saline lakes the range was between 6.76 and 36.5 × 10⁶ mL^–1. The lowest value was found in meromictic Ace Lake and the highest value in hypersaline Lake Williams. Virus to bacteria ratios (VBR) were lowest in the freshwater lakes and highest in the saline lakes, with a maximum of 23.4 in the former and 50.3 in the latter. 2. A range of morphologies among VLP was observed, including phages with short (Podoviridae) and long tails, icosahedric viruses of up to 300 nm and star‐like particles of about 80 nm diameter. 3. In these microbially dominated ecosystems there was no correlation between VLP and either bacterial numbers or chlorophyll a. There was a significant correlation between VLP abundances and dissolved organic carbon concentration (r=0.845, P < 0.01). 4. The data suggested that viruses probably attack a spectrum of bacteria and protozoan species. Virus‐like particle numbers in the freshwater lakes were lower than values reported for lower latitude systems. Those in the saline lakes were comparable with abundances reported from other Antarctic lakes, and were higher than most values published for lower latitude lakes and many marine systems. Across the salinity spectrum from freshwater through brackish to hypersaline, VLP concentrations increased roughly in relation to increasing trophy. 5. Given that Antarctic lakes have a plankton almost entirely made up of bacteria and protists, and that VLP abundances are high, it is likely that viruses play a pivotal role in carbon cycling in these extreme ecosystems.     

Paleopigment evidence of competition between phytoplankton and a cyanobacterial algal mat in a meromictic lake near Toronto,Ontario Canada

Crawford Lake, a meromictic lake located near Toronto, Canada, was cored to determine if algal pigments preserved in its sediments would make it possible to infer past changes in lake productivity over the last five hundred years. From 1500 to 1910 A.D. the sediments display extremely high levels of oscillaxanthin and myxoxanthophyll while chlorophyll derivatives and total carotenoids were relatively low. As the lake became increasingly more eutrophic in the latter part of the twentieth century, this relationship reversed itself. Competition for light between the deep dwelling cyanobacteria in the algal mat on the lake's bottom (8–14 m) and phytoplankton in the overlying surface layers of the water column (5–7 m) was attributed to the observed reduction in oscillaxanthin and myxoxanthophyll as Crawford Lake eutrophied. Because the major cyanobacteria in Crawford Lake are benthic mat forming Lyngbya and Oscillatoria, and not phytoplankton, competition for light with the overlying phytoplankton is critical in determining the total quantity of oscillaxanthin and myxoxanthophyll preserved in the lake's profundal sediments. These findings have major implications for the use of cyanobacterial pigments as indicators of lake trophic status in lakes where benthic algal mats are present.     

Microbial community structure and biomass estimates of a methanogenic Antarctic Lake ecosystem as determined by phospholipid analyses

Phospholipid analyses were performed on water column particulate and sediment samples from Ace Lake, a meromictic lake in the Vestfold Hills, Antarctica, to estimate the viable microbial biomass and community structure in the lake. In the water column, methanogenic bacterial phospholipids were present below 17 m in depth at concentrations which converted to a biomass of between 1 and 7×10⁸ cells/liter. Methanogenic biomass in the sediment ranged from 17.7×10⁹ cells/g dry weight of sediment at the surface to 0.1×10⁹ cells/g dry weight at 2 m in depth. This relatively high methanogenic biomass implies that current microbial degradation of organic carbon in Ace Lake sediments may occur at extremely slow rates. Total microbial biomass increased from 4.4×10⁸ cells/ liter at 2 m in depth to 19.4×10⁸ cells/liter at 23 m, near the bottom of the water column. Total nonarchaebacterial biomass decreased from 4.2 ×10⁹ cells/g dry weight in the surface sediment (1/4 the biomass of methanogens) to 0.06×10⁸ cells/g dry weight at 2 m in depth in the sediment. Phospholipid fatty acid profiles showed that microeukaryotes were the major microbial group present in the oxylimnion of the lake, while bacteria dominated the lower, anoxic zone. Sulfate-reducing bacteria (SRB) comprised 25% of the microbial population at 23 m in depth in the water column particulates and were present in the surface sediment but to a lesser extent. Biomass estimates and community structure of the Ace Lake eco-system are discussed in relation to previously measured metabolic rates for this and other antarctic and temperate ecosystems. This is the first instance, to our knowledge, in which the viable biomass of methanogenic and SRB have been estimated for an antarctic microbial community.     

Bacterial Communities of Three Saline Meromictic Lakes in Central Asia

Meromictic lakes located in landlocked steppes of central Asia (~2500 km inland) have unique geophysiochemical characteristics compared to other meromictic lakes. To characterize their bacteria and elucidate relationships between those bacteria and surrounding environments, water samples were collected from three saline meromictic lakes (Lakes Shira, Shunet and Oigon) in the border between Siberia and the West Mongolia, near the center of Asia. Based on in-depth tag pyrosequencing, bacterial communities were highly variable and dissimilar among lakes and between oxic and anoxic layers within individual lakes. Proteobacteria, Bacteroidetes, Cyanobacteria, Actinobacteria and Firmicutes were the most abundant phyla, whereas three genera of purple sulfur bacteria (a novel genus, Thiocapsa and Halochromatium) were predominant bacterial components in the anoxic layer of Lake Shira (~20.6% of relative abundance), Lake Shunet (~27.1%) and Lake Oigon (~9.25%), respectively. However, few known green sulfur bacteria were detected. Notably, 3.94% of all sequencing reads were classified into 19 candidate divisions, which was especially high (23.12%) in the anoxic layer of Lake Shunet. Furthermore, several hydro-parameters (temperature, pH, dissolved oxygen, H₂S and salinity) were associated (P< 0.05) with variations in dominant bacterial groups. In conclusion, based on highly variable bacterial composition in water layers or lakes, we inferred that the meromictic ecosystem was characterized by high diversity and heterogenous niches.     

Denitrification in a meromictic lake and its relevance to nitrogen flows within a moderately impacted forested catchment

We analysed the spatial and temporal variability of benthic nitrogen fluxes and denitrification rates in a sub-alpine meromictic lake (Lake Idro, Italy), and compared in-lake nitrogen retention and loss with the net anthropogenic nitrogen inputs to the watershed. We hypothesized a low nitrogen retention and denitrification capacity due to meromixis. This results from nitrate supply from the epilimnion slowing down during stratification and oxygen deficiency inhibiting nitrification and promoting ammonium recycling and its accumulation. We also hypothesized a steep vertical gradient of sedimentary denitrification capacity, decreasing with depth and oxygen deficiency. These are important and understudied issues in inland waters, as climate change and direct anthropic pressures may increase the extent of meromixis. Nearshore sediments had high denitrification rates (87 mg m^?2 day^?1) and efficiency (~ 100%), while in the monimolimnion denitrification was negligible. The littoral zone, covering 10% of the lake surface, contributed ~50% of total denitrification, while the monimolimnion, which covered 70% of the sediment surface, contributed to < 13% of total denitrification. The persistent and expanding meromixis of Lake Idro is expected to further decrease its nitrogen removal capacity (31% of the incoming nitrogen load) compared to what has been measured in other temperate lakes. Values up to 60% are generally reported for other such lakes. Results of this study are relevant as the combination of anthropogenic pressures, climate change and meromixis may threaten the nitrogen processing capacity of lakes.     

Sulfur-Oxidizing Bacteria in Soap Lake (Washington State), a Meromictic, Haloalkaline Lake with an Unprecedented High Sulfide Content

Culture-dependent and -independent techniques were used to study the diversity of chemolithoautotrophic sulfur-oxidizing bacteria in Soap Lake (Washington State), a meromictic, haloalkaline lake containing an unprecedentedly high sulfide concentration in the anoxic monimolimnion. Both approaches revealed the dominance of bacteria belonging to the genus Thioalkalimicrobium, which are common inhabitants of soda lakes. A dense population of Thioalkalimicrobium (up to 10⁷ cells/ml) was found at the chemocline, which is characterized by a steep oxygen-sulfide gradient. Twelve Thioalkalimicrobium strains exhibiting three different phenotypes were isolated in pure culture from various locations in Soap Lake. The isolates fell into two groups according to 16S rRNA gene sequence analysis. One of the groups was closely related to T. cyclicum, which was isolated from Mono Lake (California), a transiently meromictic, haloalkaline lake. The second group, consisting of four isolates, was phylogenetically and phenotypically distinct from known Thioalkalimicrobium species and unique to Soap Lake. It represented a new species, for which we suggest the name Thioalkalimicrobium microaerophilum sp. nov.     

An ecological study of two shallow,equatorial lakes: Lake Mburo and Lake Kachera,Uganda

Lake Mburo and Lake Kachera are shallow, eutrophic lakes in mid‐western Uganda. Lake Mburo recorded higher values of Secchi and eutrophic depths and lower extinction coefficient (k) values. The lakes showed a ‘red shift’ phenomenon in maximum light transmission. The average values of electrical conductivity in Lake Mburo and Lake Kachera were 136 and 244 μS cm^?1, respectively. The pH values indicated high photosynthetic activity. Dissolved oxygen concentration averaged 6.9 and 7.8 mg l^?1 in Lake Mburo and Lake Kachera, respectively. The lakes had high total nitrogen (TN) : total phosphorus (TP) ratios averaging 200 and 280 in Lake Mburo and Lake Kachera, respectively. The lakes are dominated by cyanobacterial blooms that reduce light penetration to less than 1 m. Lake Mburo had a lower algal biomass than Lake Kachera. Chlorophyll a concentrations correlated positively (r = 0.73, P < 0.05) with the extinction coefficient in Lake Mburo but not in Lake Kachera. The correlations between chlorophyll a and TN and TP were also high. Both lakes recorded high primary productivity, Lake Mburo showing higher values. The study highlighted the need to investigate the organism–community interrelationships in the two water bodies.     

Plankton communities of three Central Florida lakes

A study of planktonic primary productivity and community structure was carried out on three lakes of varying morphometric and chemical features, but which were within a single watershed in Central Florida. Primary productivity in these lakes was evaluated by means of in situ light and dark bottle determinations as well as by calculations based upon chlorophyll and light data. Lake Mize, a deep sinkhole lake containing highly colored waters of low pH, proved to be a very unproductive lake, while Biven's Arm, a shallow-basin lake with alkaline waters and Newnan's lake, a shallow-basin like with highly colored waters both showed relatively high productivities. Phytoplankton components of the two productive lakes were quite similar to one another in being composed of bloom-forming blue-green and green algae, while the predominant forms in Lake Mize were chrysophytes, dinoflagellates and a diatom. Zooplanktonic forms were more uniform among the lakes than were the phytoplankton, though population levels reflected relative productivities among the lakes. Rotifers predominated in the Lake Mize plankton while cladocerans were relatively inabundant. This was opposite to the condition in Biven's Arm and Newnan's Lake. It is suggested that the constant presence of larval Chaoborus in the water column in Lake Mize may be partially responsible for the paucity of cladocera.     

Aquatic macrophytes in saline lakes of the Canadian prairies

Vascular macrophyte species richness decreases with increasing salinity. Only three species of submerged plants (Potamogeton pectinatus, Ruppia maritima, R. occidentalis) tolerate hypersaline waters (>50 g l^-1, total of ionic constituents). Eight emergent species occur in more saline habitats but only five (Scirpus maritimus var. paludosus, Distichlisstricta, Puccinellia nuttalliana, Scirpus americanus, Triglochin maritima) occur commonly over a range of saline lakes into the hypersaline category. Usually, species tolerant of high salinities are found over the entire saline spectrum and even extend into subsaline waters (<3 g l^-1) and thrive there. A major increase in the number of species occurs below 5 g l^-1. As the water recedes plants such as Salicornia rubra, Suaeda calceoliformes, Hordeum jubatum and Sonchus arvensis invade.Submerged angiosperm distribution is controlled by total ion concentration and substrate texture plays no apparent role. Although angiosperms normally grow in all kinds of substrates, they occupy coarse substrates in Wakaw lake because suitable fine substrates are densely colonized by charophytes. In this lake light limited growth occurs to a depth of 5% of surface light. Light was not limiting in Redberry Lake but angiosperm growth was limited to the upper 8 m (10% or more of surface light). Thermal stratification and depth (pressure) were probably limiting istead. In meromictic Waldsea Lake the depth of the chemocline (6 m, 5% surface light) delimits angiosperm growth.     

The microbial composition of three limnologically disparate hypersaline Antarctic lakes

16S rRNA clone library analysis was used to examine the biodiversity and community structure within the sediments of three hypersaline Antarctic lakes. Compared to sediment of low to moderate salinity Antarctic lakes the species richness of the hypersaline lake sediments was 2-20 times lower. The community of Deep Lake (32% salinity, average sediment temperature -15 degrees C) was made up almost entirely of halophilic Archaea. The sediment communities of two meromictic hypersaline lakes, Organic Lake (20% salinity, -7 degrees C) and Ekho Lake (15% salinity, 15 degrees C) were more complex, containing phylotypes clustering within the Proteobacteria and Cytophagales divisions and with algal chloroplasts. Many phylotypes of these lakes were related to taxa more adapted to marine-like salinity and perhaps derive from bacteria exported into the sediment from the lower salinity surface waters. The Ekho Lake clone library contained several major phylotypes related to the Haloanaerobiales, the growth of which appears to be promoted by the comparatively high in situ temperature of this lake.     

Grazing rates of a calanoid copepod (Paralabidocera antarctica) in a continental Antarctic lake

The presence of crustaceans in some Antarctic lakes adds a level of complexity to the food webs of these environments. The grazing pressure of the crustaceans on phytoplankton and other protists, and the role they play in controlling food web structure, has not yet been examined in any detail. This paper presents the first data on grazing pressure of crustacean zooplankton in continental Antarctic lakes. In this study the grazing rates of Paralabidocera antarctica in saline Ace Lake, Vestfold Hills, were assessed using a radio-tracer method. Clearance rates ranged from approximately 1 to 8 ml ind.⁻¹ day⁻¹, depending on developmental stage. Due to their high abundance, nauplii exhibited the highest daily carbon ingestion rates. The population cleared about 3% of the water column per day, accounting for a significant fraction of primary production. P. antarctica has a considerable impact on the carbon cycle in Ace Lake and, by inference, in the other lakes in which it occurs. Accepted: 30 October 1999     

The vertical distribution and abundance of <Emphasis Type="Italic">Gammarus lacustris</Emphasis> in the pelagic zone of the meromictic lakes Shira and Shunet (Khakassia,Russia)

The vertical distribution and abundance of Gammarus lacustris in the pelagic zone of two fishless meromictic lakes, L. Shira and L. Shunet, in Southern Siberia (Russia), was studied with the underwater video recording system and using vertical hauls. In both lakes, during summer stratification, Gammarus was distributed non-homogenously, with a stable peak in the metalimnion. The average depth of Gammarus population in the pelagic zone was significantly correlated with the depth of the thermocline. Gammarus abundances obtained using vertical plankton hauls with net were quite comparable with those obtained from video records. The peak abundance of Gammarus in the pelagic zone of the lakes observed with underwater video amounted up to 400 individuals m⁻², while the peak animal densities in the metalimnion reached 50 ind. m⁻³. The data are compared with previously published abundances of Gammarus in the littoral of Lake Shira. Both littoral and pelagic can be equally important habitats for amphipods in meromictic lakes. The absence of fish in the pelagic zone, high oxygen concentration, low water temperature, increased seston concentration, elevated water density in the metalimnion and the anoxic hypolimnion can be the most probable combination of factors that are responsible for the peak of Gammarus in the metalimnion of these lakes.