Limnology....The Long Version
Little
is known about limnological conditions in subglacial lakes. Knowledge of
physical, chemical, and microbiological processes within “lakes” will be
essential for detecting the presence of life, understanding the associated
biogeochemistry and deciphering sedimentary records of lake histories. There is
a range in lake sizes, morphologies, and geological settings that reflects a
spectrum of ages, environmental conditions, and habitats. Model predictions of
circulation and analyses of accreted lake water have been used to speculate
about subglacial lake limnology in general and Lake Vostok’s
chemical environment in particular.
Physical
models assessing the likelihood of water circulation in Lake Vostok
have been developed by Wüest and Carmack (2000), Williams (2001) and Mayer et
al. (2003) and recently reviewed by Siegert (2005). Lake
Vostok circulation is predicted to be
a consequence of differences in the pressure melting point between the north
and south ends of Lake
Vostok. In a freshwater
(i.e. low ionic strength) lake, geothermal heating warms bottom waters to a
temperature higher than that of the upper layers. Water density decreases with
increasing temperature resulting in an unstable water column leading to
convective circulation where cold melt water sinks and water warmed by
geothermal heat ascends (Wüest & Carmack 2000). Conversely, if the lake is
slightly saline the fresh glacier melt water will be buoyant relative to bulk
lake water and the northern melt water would spread southward and upward
(Souchez et al. 2000). If the horizontal salinity gradient is great enough to
compensate for geothermal warming, water would move into regions of
progressively lower pressure displacing lake water in the south. The cold
northern water eventually refreezes onto the ice sheet base some distance from
where it first melted. In this case, a conveyor of fresh cool melt water
migrates from north to south beneath the ice sheet causing displacement of
warmer dense lake water from the south to the north. If the lakes exhibit
vertical chemical gradients a stratified upper layer of cold fresh water would
be predicted to overlie a deeper layer of warm saline water.
Several
authors have discussed the geochemistry of Lake Vostok
as inferred from the analysis of proxies in accreted lake ice. Accreted ice is
ice frozen onto the base of the ice sheet as it moves across the lake surface.
As described above, this process is the result of a differential in the
thickness of the overlying ice that establishes a melt/freeze cycle within a
lake basin. The freezing process imparts distinct stable oxygen and hydrogen
isotopic compositions to accretion ice water that is significantly different
than the overlying glacial ice (Jouzel et al, 1999; Souchez et al, 2003). Two
types of accretion ice have been recovered from the Vostok borehole: (i) ice
with sediment/particle inclusions and (ii) ice with no visible inclusions (e.g.
Jouzel et al, 1999, Souchez et al, 2003). It is thought that the ice with no
visible inclusions is water from the deeper portions of the lake while the ice
containing visible inclusions may have entrained material as it intersected the
bedrock upstream of Vostok Station (Jouzel et al, 1999). Smaller inclusions
consist of mostly fine clays, quartz aggregates and various salts. Dating
suggests a Precambrian origin for the inclusions consistent with a continental
shield source (Delmonte et al, 2004).
Gas
concentrations and isotopic ratios in accreted ice lend clues to the origins of
Lake Vostok water. Helium isotope ratios in
accretion ice are lower than glacier ice (Jean Baptiste et al, 2001) suggesting
an input of radiogenic helium from the degassing of deep faults (Bulat et al,
2004). In contrast, 3He
concentrations suggest little if any hydrothermal input to Lake Vostok
(Jean Baptiste et al, 2001). Seismic activity near Vostok station may
contribute heat and chemicals to the lake but this conjecture remains
speculative (Studinger et al, 2003; Bulat et al, 2004).
Gas
hydrate is suspected of playing a role in establishing the physical, chemical,
and biological characteristics of subglacial lakes. Atmospheric air, captured in
ice sheets, occurs exclusively as gas hydrate at an ice thickness of a few
kilometers (Hondoh, 1996; Uchida etal, 1994). In large subglacial lakes, such
as Lake Vostok, with a geometry that favors the
establishment of a melt/freeze cycle, the melting of the ice sheet would
release air hydrate to the water. Accretion ice is nearly gas-free relative to
overlying glacial ice due to the exclusion of gas and salt during refreezing
(Jouzel et al. 1999). This exclusion would lead to increased dissolved gas concentrations
in Lake Vostok (McKay et al. 2003). Dissolved
oxygen concentrations are predicted to be many times higher than
air-equilibrated water (McKay et al. 2003). Within 400,000 years, gas
concentrations in the lake are predicted to reach levels that favor the
formation of gas hydrate. Gas hydrate formed in the lake would float to the
surface of the lake if formed from air or sink if it contained significant CO2.
These estimates of Lake
Vostok water dissolved
oxygen concentrations do not consider the effects of removal processes (McKay
et al. 2003; Lipenkov et al. 2001). Metabolic consumption of oxygen in bottom
waters and/or sediments can reduce oxygen levels and balance the accumulation
of gases due to gas hydrate disassociation (e.g., McKay et al. 2003).
Dissolved
organic carbon (DOC) plays a key role in ecosystem carbon cycling owing to its
role as an energy and carbon source for heterotrophic organisms. DOC levels in Lake Vostok
accretion ice are low implying that DOC levels in surface lake waters are low
as well. However, little is known about the partitioning coefficients of DOC as
water freezes so the actual DOC levels in the lake remain unknown (for further
details see the next section on Microbial Life, Evolution, and Adaptations).
A
more complete description of the limnology and biogeochemical processes at work
in subglacial environments is fundamental to understanding subglacial
environments as systems. Scientific questions related to limnology and
biogeochemistry will be answered with a combination of models, field
observations, in situ experiments and
monitoring, and laboratory measurements (see Inset). Refined models of
circulation will require detailed knowledge of basin topography and water
density. The equations of state used to compute density from salinity and
temperature data in seawater may not apply to subglacial lakes if the ion
ratios differ significantly from seawater. It will be important to test the
models using natural tracers (e.g. isotopic compositions of Be, He, rare earth
elements, etc). Given the potential freeze/thaw nature of some of these
systems, it will also be important to understand the dynamics of salt rejection
and partitioning that occurs at the ice-water interface. The potentially low
kinetic energy of these systems in concert with steep-sided basins may lead to
vertical density stratification that inhibits mixing of near-surface and
near-bottom lake waters. In such cases, thermohaline convection may be an
important vertical transport mechanism. In order to understand the density
structure of the lakes, detailed profiles of the ionic content of the water
column are needed. If there is a geothermal source of heat and chemicals in
some subglacial environments, it will play an important role in establishing
buoyancy driven circulation and determining the ionic composition of lake
waters. Critical to understanding subglacial lake circulation patterns is the
possibility of episodic or continuous hydraulic connectivity between lake
systems. Outbursts of subglacial melt waters would “reset” water density
gradients homogenizing the physical, chemical, and biological environment. It
is critically important to know if these environments are “open” or “closed”
systems, i.e., is there exchange among lakes within a certain geographical area
and on what timeframes. To model circulation; construct water, heat, and
biochemical budgets; and estimate the age of the water, the basin topography, salinity,
temperature, density, and current distributions must be accurately known.
Biogeochemical studies will provide important data documenting the presence and type of metabolic activity. Metabolic reactions require a number of essential elements. The amount and distribution of nutritive elements (e.g., nitrogen, phosphorus, sulfur, iron and others) in subglacial environments will provide fundamental constraints on biological productivity and suggest how microorganisms meet their metabolic needs. It will be essential to document the spatial and temporal distribution of various elements in the water column and sediments to understand their cycling. Information on potentially toxic elements (i.e., oxygen radicals) will lend clues to the habitability of subglacial environments. Characterization of the water column will include profiles of salinity, temperature, density, dissolved O2, the CO2 system, pH, Eh, and characterization of dissolved and particulate organic matter. Observatories with strings of sensors will be deployed in water columns to detect temporal changes in water properties. Discrete samples taken in profile within the water column will be analyzed for biogenic gases such as N2O, H2S, N2, and CH4; inorganic N (NH4+, NO2-, N2O); iron; soluble reactive phosphorus; dissolved organic carbon; dissolved inorganic carbon; major cations and anions; particulate carbon and nitrogen; microorganism; and 16S and 18S r-RNA gene microbial diversity. Eventually, these same measurements will be performed on sediment samples, including vertical profiles, within sedimentary columns to establish the presence of chemical gradients that support biological activity.

