Perennially ice-covered Lake Hoare,Antarctica: physical environment,biology and sedimentation

Lake Hoare (77° 38 S, 162° 53 E) is a perennially ice-covered lake at the eastern end of Taylor Valley in southern Victoria Land, Antarctica. The environment of this lake is controlled by the relatively thick ice cover (3–5 m) which eliminates wind generated currents, restricts gas exchange and sediment deposition, and reduces light penetration. The ice cover is in turn largely controlled by the extreme seasonality of Antarctica and local climate. Lake Hoare and other dry valley lakes may be sensitive indicators of short term (< 100 yr) climatic and/or anthropogenic changes in the dry valleys since the onset of intensive exploration over 30 years ago. The time constants for turnover of the water column and lake ice are 50 and 10 years, respectively. The turnover time for atmospheric gases in the lake is 30–60 years. Therefore, the lake environment responds to changes on a 10–100 year timescale. Because the ice cover has a controlling influence on the lake (e.g. light penetration, gas content of water, and sediment deposition), it is probable that small changes in ice ablation, sediment loading on the ice cover, or glacial meltwater (or groundwater) inflow will affect ice cover dynamics and will have a major impact on the lake environment and biota.     

Temporal plankton dynamics in an oligotrophic maritime Antarctic lake

• 1 The population density, diversity and productivity of the microbial plankton in an oligotrophic maritime Antarctic lake were studied for a 15‐month period between December 1994 and February 1996.
• 2 In the lake, concentrations of nutrients and dissolved organic carbon were uniformly low, temperature varied over a small annual range of 0.1–3 °C, and the surface was ice‐covered except during a period of approximately 6 weeks in summer.
• 3 The total of 57 morphotypes of protozoa observed during the study is a higher taxonomic diversity than previously reported from continental Antarctic lakes, but lower than that found in more eutrophic maritime Antarctic lakes. Likewise, planktonic abundance and productivity were lower than has been reported in other lakes on Signy Island, but generally higher than those of lakes on the Antarctic continent.
• 4 There were marked seasonal and interannual variations in planktonic population density.
• 5 Chlorophyll a concentrations ranged from undetectable to 4.2 µg L^‐1 and the greatest rate of primary productivity measured was 4.5 mg C m^‐3 h^‐1. The phytoplankton was dominated by small chlorophytes and chrysophytes, with phototrophic nanoflagellate abundance ranging from 1.1 × 10³ to 1.2 × 10⁷ L^‐1.
• 6 Bacterial densities of 3.6 × 10⁸ to 1.9 × 10¹⁰ L^‐1 were recorded and bacterial productivity reached a peak of 0.36 µg C L^‐1 h^‐1. Numbers of heterotrophic nanoflagellates between 5.0 × 10⁴ and 1.8 × 10⁷ L^‐1, and of ciliates from undetectable to 1.1 × 10⁴ L^‐1 were observed. Naked amoebae were usually rare, but occasionally reached peaks of up to 1.5 × 10³ L^‐1.

     

Annual growth layers as proxies of past growth conditions for benthic microbial mats in a perennially ice-covered Antarctic lake

Perennial microbial mats can be the dominant autotrophic community in Antarctic lakes. Their seasonal growth results in clearly discernible annual growth layering. We examined features of live microbial mats from a range of depths in Lake Hoare, Antarctica, that are likely to be preserved in these layers to determine their potential as proxies of past growth performance. Cyanobacteria dominated the mat for all but the deepest depth sampled. Changes in areal concentrations of phycobilin pigments, organic matter and extracellular polysaccharide and in species composition did not correspond to changes in various water column properties, but showed a linear relationship with irradiance. Carbonate accumulation in the mats correlated with biomass markers and may be inferred as an index of mat performance. We examined the carbonate content of annual layers laid down from 1958–1959 to 1994–1995 in sediment cores from 12 m depth. The carbonate content in the layer showed a significant correlation with the mean summer air temperature. These data suggest a link between air temperature and microbial mat growth performance, and suggest that it is mediated via irradiance. Laminated microbial mats in Antarctic lakes have the potential to act as fine-resolution records of environmental conditions in the recent past, although interpretation is complex.     

Legacies of recent environmental change in the benthic communities of Lake Joyce, a perennially ice-covered Antarctic lake

Many Antarctic lakes provide habitat for extensive microbial mats that respond on various timescales to environmental change. Lake Joyce contains calcifying microbialites and provides a natural laboratory to constrain how environmental changes influence microbialite development. In Lake Joyce, depth-specific distributions of calcitic microbialites, organic carbon, photosynthetic pigments and photosynthetic potential cannot be explained by current growth conditions, but are a legacy of a 7-m lake level rise between 1973 and 2009. In the well-illuminated margins of the lake, photosynthetically active benthic communities colonised surfaces submerged for just a few years. However, observed increases in accumulated organic material with depth from 5 to 20 m (2-40 mg ash-free dry weight cm(-2)) and the presence of decimetre-scale calcite microbialites at 20-22 m depth, apparently related to in situ photosynthetic growth, are inconsistent with the current distributions of irradiance, photosynthetic pigments and mat photosynthetic potential (as revealed by pulse-amplitude-modulated fluorometry). The microbialites appeared photosynthetically active in 1986 and 1997, but were outside the depth zone where significant phototrophic growth was possible and were weakly photosynthetically competent in 2009. These complex microbial structures have persisted after growth has ceased, demonstrating how fluctuating environmental conditions and the hysteresis between environmental change, biological response and microbialite development can be important factors to consider when interpreting modern, and by inference ancient, microbially mediated structures.     

Cold-active acetogenic bacteria from surficial sediments of perennially ice-covered Lake Fryxell, Antarctica

RibR is a minor cryptic flavokinase (EC 2.7.1.26) of the Gram-positive bacterium Bacillus subtilis with an unknown cellular function. The flavokinase activity appears to be localized to the N-terminal domain of the protein. Using the yeast three-hybrid system, it was shown that RibR specifically interacts in vivo with the nontranslated wild-type leader of the mRNA of the riboflavin biosynthetic operon. This interaction is lost partially when a leader containing known cis-acting deregulatory mutations in the so-called RFN element is tested. The RFN element is a sequence within the rib-leader mRNA reported to serve as a receptor for an FMN-dependent 'riboswitch'. In RibR itself, interaction was localized to the carboxy-terminate part of the protein, a segment of unknown function that does not show similarity to other proteins in the public databases. Analysis of a ribR-defective strain revealed a mild deregulation with respect to flavin (riboflavin, FMN and FAD) biosynthesis. The results indicate that the RNA-binding protein RibR may be involved in the regulation of the rib genes.     

Evolution of Antarctic lake ecosystems

Antarctic lakes present a wide variety of physical, chemical and biological conditions, and are not always the simplified systems imagined by earlier workers. The volume of data on lakes of various ages now allows informed speculation on the evolution of the Antarctic lake ecosystem.     

Evolution of temperature and salt structure of Lake Bonney, a chemically stratified Antarctic lake

A resurgence of interest in the ecology of perennially ice-covered lakes in the McMurdo dry valleys has necessitated a review of our knowledge of the physical and chemical properties of these unusual lakes. Salinities in the ice-covered lakes cover a range from freshwater to hypersaline brines. Recent measurements of salt composition and concentrations in Lake Bonney reveal little change below the chemocline since extensive measurements made in 1960–1961, although lake level has risen by approximately 5 m since that time. The rise in lake level has resulted in a thickening of the freshwater layer above the chemocline. Temperature structure has adjusted to the effects of increased lake level on heat transfer processes such as transmission and absorption of solar radiation in the water column.Questions about how water-column stability affects biology in Lake Bonney have motivated the formulation of a method to compute density from in situ measurements of temperature, conductivity and pressure. Owing to high salt concentration and unique ion ratios, we modified the UNESCO Equation of State for seawater to predict density at salinities greater than 42. The modifications merge smoothly with the UNESCO equations at a salinity of 42. At salinities below 42 the UNESCO equations give excellent predictions of density.     

Factors regulating summer phytoplankton in a highly eutrophic Antarctic lake

Lakes from Maritime Antarctica are regarded as systems generally inhabited by metazoan plankton capable of imposing a top-down control on the phytoplankton during short periods, while lakes from Continental Antarctica lacking these communities would be typically controlled by scarcity of nutrients, following a bottom-up model. Otero Lake is a highly eutrophic small lake located on the NW of the Antarctic Peninsula, which has no metazoan plankton. During summer 1996, we studied the density, composition and vertical distribution of the phytoplankton community of this lake with respect to various abiotic variables, yet our results demonstrated neither light nor nutrient limitation of the phytoplankton biomass. Densities of heterotrophic nanoflagellates (HNAN) and ciliates from three different size categories were also studied. Extremely low densities of HNAN (0–155 ind. ml^–1) could be due to feeding competition by bacterivore nanociliates and/or predation by large ciliates. A summer bloom of the phytoflagellate Chlamydomonas aff. celerrima Pascher reached densities tenfold those of previous years (158.10³ ind. ml^–1), though apparently curtailed by a strong peak of large ciliates (107 ind. ml^–1) which would heavily graze on PNAN (phototrophic nanoflagellates). Top-down control can thus occur in this lake during short periods of long hydrologic residence time.     

Exploration of Ellsworth Subglacial Lake: a concept paper on the development, organisation and execution of an experiment to explore, measure and sample the environment of a West Antarctic subglacial lake

Antarctic subglacial lakes have, over the past few years, been hypothesised to house unique forms of life and hold detailed sedimentary records of past climate change. Testing this hypothesis requires in situ examinations. The direct measurement of subglacial lakes has been considered ever since the largest and best-known lake, named Lake Vostok, was identified as having a deep water-column. The Subglacial Antarctic Lake Environments (SALE) programme, set up by the Scientific Committee on Antarctic Research (SCAR) to oversee subglacial lakes research, state that prior exploration of smaller lakes would be a “prudent way forward”. Over 145 subglacial lakes are known to exist in Antarctica, but one lake in West Antarctica, officially named Ellsworth Subglacial Lake (referred to hereafter as Lake Ellsworth), stands out as a candidate for early exploration. A consortium of over 20 scientists from seven countries and 14 institutions has been assembled to plan the exploration of Lake Ellsworth. An eight-year programme is envisaged: 3 years for a geophysical survey, 2 years for equipment development and testing, 1 year for field planning and operation, and 2 years for sample analysis and data interpretation. The science experiment is simple in concept but complex in execution. Lake Ellsworth will be accessed using hot water drilling. Once lake access is achieved, a probe will be lowered down the borehole and into the lake. The probe will contain a series of instruments to measure biological, chemical and physical characteristics of the lake water and sediments, and will utilise a tether to the ice surface through which power, communication and data will be transmitted. The probe will pass through the water column to the lake floor. The probe will then be pulled up and out of the lake, measuring its environment continually as this is done. Once at the ice surface, any water samples collected will be taken from the probe for laboratory analysis (to take place over subsequent years). The duration of the science mission, from deployment of the probe to its retrieval, is likely to take between 24 and 36 h. Measurements to be taken by the probe will provide data about the following: depth, pressure, conductivity and temperature; pH levels; biomolecules (using life marker chips); anions (using a chemical analyzer); visualisation of the environment (using cameras and light sources); dissolved gases (using chromatography); and morphology of the lake floor and sediment structures (using sonar). After the probe has been retrieved, a sediment corer may be dropped into the lake to recover material from the lake floor. Finally, if time permits, a thermistor string may be left in the lake water to take time-dependent measurements of the lake’s water column over subsequent years. Given that the comprehensive geophysical survey of the lake will take place in two seasons during 2007–2009, a two-year instrument and logistic development phase from 2008 (after the lake’s bathymetry has been assessed) makes it possible that the exploration of Lake Ellsworth could take place at the beginning of the next decade.     

Mixotrophic cryptophytes and their predators in the Dry Valley lakes of Antarctica

1. The ingestion rates of planktonic, mixotrophic cryptophytes in two perennially ice-covered Antarctic lakes in the McMurdo Dry Valleys, were investigated during the summer of 1997–1998.
2. In Lake Fryxell, which is meromictic, ingestion rates increased with depth in November and were highest in a cryptophyte maximum close to the chemocline. In Lake Hoare, which is unstratified and freshwater, there was no significant difference in ingestion rates with depth. In both lakes, the highest ingestion rates occurred in early summer, decreasing in December and January. Ingestion rates varied between 0.2 bacteria cell⁻¹ h⁻¹ and 3.6 bacteria cell⁻¹ h⁻¹.
3. During November, mixotrophic cryptophytes removed up to 13% of bacterial biomass day⁻¹ and had a greater grazing impact than heterotrophic nanoflagellates (HNAN). As summer progressed, the grazing impact of cryptophytes and HNAN became similar.
4. The maximum depth of cryptophytes in Lake Fryxell was predated by a population of the ciliate Plagiocampa. Plagiocampa had an ingestion rate of 0.13–0.19 cryptophytes cell⁻¹ h⁻¹. The grazing impact on the cryptophyte community was insignificant. However, the ciliate appeared to be indulging in temporary mixotrophy, sequestering the cryptophytes for a number of weeks before digesting them.
5. It is suggested that mixotrophy is an important survival strategy in the extreme lake ecosystems of the McMurdo Dry Valleys.     

Alteration of the food web along the Antarctic Peninsula in response to a regional warming trend

In the nearshore coastal waters along the Antarctic Peninsula, a recurrent shift in phytoplankton community structure, from diatoms to cryptophytes, has been documented. The shift was observed in consecutive years (1991–1996) during the austral summer and was correlated in time and space with glacial melt‐water runoff and reduced surface water salinities. Elevated temperatures along the Peninsula will increase the extent of coastal melt‐water zones and the seasonal prevalence of cryptophytes. This is significant because a change from diatoms to cryptophytes represents a marked shift in the size distribution of the phytoplankton community, which will, in turn, impact the zooplankton assemblage. Cryptophytes, because of their small size, are not grazed efficiently by Antarctic krill, a keystone species in the food web. An increase in the abundance and relative proportion of cryptophytes in coastal waters along the Peninsula will likely cause a shift in the spatial distribution of krill and may allow also for the rapid asexual proliferation of carbon poor gelatinous zooplankton, salps in particular. This scenario may account for the reported increase in the frequency of occurrence and abundance of large swarms of salps within the region. Salps are not a preferred food source for organisms that occupy higher trophic levels in the food web, specifically penguins and seals, and thus negative feedbacks to the ecology of these consumers can be anticipated as a consequence of shifts in phytoplankton community composition.     

Phytoplankton structure and dynamics in a volcanic lake in Deception Island (South Shetland Islands,Antarctica)

This work constitutes the first floristic and ecological analysis of the phytoplankton community of a volcanic freshwater lake in Deception Island (62°57′S, 60°38′W, South Shetland Islands, Antarctica). The main limnological features and phytoplankton size fractions were analyzed. Samples were taken during the austral summer of 2002 at two opposite sites. According to ANOVA results performed with abiotic variables, no significant differences between sites were found. The phytoplankton community showed low algal species richness, with an important contribution of the tychoplanktonic taxa. In terms of species number, Bacillariophyceae was the dominant class. Autotrophic picoplankton registered the highest densities from the second sampling date onwards. Nanophytoplankton was represented by unidentified chrysophycean organisms, which showed different distribution patterns between sites. The net phytoplankton abundance remained low during the sampling period and was strongly correlated with chlorophyll a concentration. Both nutrient concentrations and chlorophyll a values indicated oligotrophic conditions.     

Plankton dynamics in a high mountain lake (Las Yeguas,Sierra Nevada,Spain). Indirect evidence of ciliates as food source for zooplankton

A detailed sampling programme during the ice-free season (July–September) in the oligotrophic lake Las Yeguas (Southern Spain) has shown a well-defined time lag between phytoplankton and zooplankton maximum standing stocks, the former displaying a peak (23 μgC l^-1) just after the ice-melting, and the latter by the end of September (80 μgC l^-1). A ratio of autotrophs to heterotrophs lower than 1 which lasted more than two thirds of the study period may suggest a high algal productivity per unit of biomass. The estimated strong top-down regulation of phytoplankton by zooplankton indicates an efficient utilization of resources. A comparative analysis between the available food supply and the critical food concentration that is necessary to maintain the population of Daphnia pulicaria (which constitutes up to 98% of the heterotrophic biomass) proves this species to be food-limited in the lake under study. To explain the dominance (and development) of such large-bodied cladoceran population, we discuss the possibility of the utilization of naked protozoan ciliates (Oligotrichidae) as a complementary high quality food source, and the exploitation of benthic resources through a coupled daily migration behaviour.     

Protist diversity in a permanently ice-covered Antarctic Lake during the polar night transition

The McMurdo Dry Valleys of Antarctica harbor numerous permanently ice-covered lakes, which provide a year-round oasis for microbial life. Microbial eukaryotes in these lakes occupy a variety of trophic levels within the simple aquatic food web ranging from primary producers to tertiary predators. Here, we report the first molecular study to describe the vertical distribution of the eukaryotic community residing in the photic zone of the east lobe (ELB) and west lobe (WLB) of the chemically stratified Lake Bonney. The 18S ribosomal RNA (rRNA) libraries revealed vertically stratified populations dominated by photosynthetic protists, with a cryptophyte dominating shallow populations (ELB–6 m; WLB–10 m), a haptophyte occupying mid-depths (both lobes 13 m) and chlorophytes residing in the deepest layers (ELB–18 and 20 m; WLB–15 and 20 m) of the photic zone. A previously undetected stramenopile occurred throughout the water column of both lobes. Temporal variation in the eukaryotic populations was examined during the transition from Antarctic summer (24-h sunlight) to polar night (complete dark). Protist diversity was similar between the two lobes of Lake Bonney due to exchange between the photic zones of the two basins via a narrow bedrock sill. However, vertical and temporal variation in protist distribution occurred, indicating the influence of the unique water chemistry on the biology of the two dry valley watersheds.     

Detection and characterization of denitrifying bacteria from a permanently ice-covered Antarctic Lake

Denitrifying bacterial strains were isolated from Lake Bonney,apermanently ice-covered and chemically stratified lake in theMcMurdo dry valley region of Antarctica, using complex mediaat4 °C. Three strains, identified as denitrifiers bytheirability to produce nitrous oxide using nitrate or nitrite as arespiratory substrate, were characterized as to theirtemperatureand salinity optima for aerobic growth in batch culture; allthreewere psychrophilic and moderately halophilic. Maximum growthratesof near 0.024 h^–1 were measured for all three strains.Growthrates projected to occur at in situ temperature andsalinityimply generation times on the order of 100 h. Species specificpolyclonal antisera were prepared against two of the strains,ELB17 (from the east lobe of the lake at 17 m) and WLB20 (fromthewest lobe at 20 m). Both strains were subsequently detectedandenumerated in the lake using the antisera. ELB17 was presentinboth lobes below the chemocline, while WLB20 was present inthewest lobe below the chemocline but only in surface waters oftheeast lobe. These distributions are related to the observedchemicaldistributions which imply the occurrence of denitrification inthewest lobe of the lake and not in the east lobe.     

Global soil ciliate (Protozoa, Ciliophora) diversity: a probability-based approach using large sample collections from Africa, Australia and Antarctica

Large sample collections from Africa (92 samples), Australia (157) and Antarctica (90) were investigated for soil ciliates using the non-flooded Petri dish method, which re-activates the ciliates' resting cysts from air-dried samples. Species were determined from life and by silver impregnation. The African samples were the richest, containing 507 species (240 undescribed,=47%), followed by the Australian (361 species, 154=43% undescribed) and the Antarctic (95 species, 14=15% undescribed) samples. The percentage of new species/sample was consistently low, viz. 4–8% on average, indicating that new species were considerably undersampled relative to described ones, very probably due to methodological shortcomings, i.e. usually only cysts of the more euryoecious species could be reactivated. Thus, a probability theory-based statistical approach was applied to the data sets to compensate for the underestimated number of undescribed species. This procedure indicated that, depending on the region, 70–80% of the soil ciliates are still unknown and global soil ciliate diversity amounts to at least 1330–2000 species. Several indicators, especially the constant rate at which new species have been found during a 20-year period of intensive research, suggest that this estimate is conservative. This revised version was published online in August 2006 with corrections to the Cover Date.     

Sixty years of environmental change in the world's largest freshwater lake – Lake Baikal, Siberia

High-resolution data collected over the past 60 years by a single family of Siberian scientists on Lake Baikal reveal significant warming of surface waters and long-term changes in the basal food web of the world's largest, most ancient lake. Attaining depths over 1.6 km, Lake Baikal is the deepest and most voluminous of the world's great lakes. Increases in average water temperature (1.21 °C since 1946), chlorophyll a (300% since 1979), and an influential group of zooplankton grazers (335% increase in cladocerans since 1946) may have important implications for nutrient cycling and food web dynamics. Results from multivariate autoregressive (MAR) modeling suggest that cladocerans increased strongly in response to temperature but not to algal biomass, and cladocerans depressed some algal resources without observable fertilization effects. Changes in Lake Baikal are particularly significant as an integrated signal of long-term regional warming, because this lake is expected to be among those most resistant to climate change due to its tremendous volume. These findings highlight the importance of accessible, long-term monitoring data for understanding ecosystem response to large-scale stressors such as climate change.     

Primary producer dynamics associated with evaporative concentration in a shallow,equatorial soda lake (Lake Elmenteita,Kenya)

Lake Elmenteita (0°27S, 36°15E) lies on the floor of the rift valley at 1776 m above sea level in Kenya. As a consequence of lower than average rainfall, the mean depth decreased from 1.1 to 0.65 m during the study period (February 1973 to August 1974). The initiation of major biological changes coincided with a period of rapid evaporative concentration in 1973 (February to April) when the conductivity increased from 19.1 to 27.0 mmhmos cm^-1. Spirulina platensis, Spirulina laxissima and Anabaenopsis arnoldii decreased in abundance precipitously in parallel with large declines in chlorophyll a concentration and phytoplankton photosynthetic rates. Once the overall abundance of phytoplankton had declined and the transparency had increased, primary productivity by benthic algae increased significantly. Paradiaptomus africanus, the only copepod living in the lake, was abundant in February and March 1973, but was gone by May. Eight hypotheses to explain these changes are evaluated and converge on the suggestion that a rate of change of salinity greater than 5 mmhmos cm^-1 per month and a salinity exceeding 25 mmhmos cm^-1 cannot be tolerated by P. africanus and adversely effects the nitrogen fixer, A. arnoldii. Furthermore, the loss of P. africanus and oxygenation of the sediments by benthic algae reduce the rate of recyling of nutrients which alters phytoplankton abundance and species composition.