Under‐ice metabolism in a shallow lake in a cold and arid climate

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Production and hydrochemical characteristics of ice,under-ice water and sediments in the Razdolnaya River estuary (Amursky Bay,Sea of Japan) during the ice cover period

Production and hydrochemical characteristics of ice, under-ice water, and sediments in the Razdol’naya River estuary (Sea of Japan) were studied during the ice cover periods of the years 2007 and 2008. In 2007, snow cover was absent until mid-February and PAR levels under ice were sufficient for the development of phytoplankton. The chlorophyll a content in ice, under-ice water, and surface sediments was high, while nutrient levels were decreased. After a snowfall, the chlorophyll content in ice and under-ice water decreased sharply. In winter 2008, snow cover was formed immediately after freeze-up; therefore, PAR levels in the ice and under-ice water were significantly reduced. The chlorophyll content was lower, but nutrient levels were higher than in 2007. In both winter seasons, the greatest portion of chlorophyll (up to 85%) was contained in surface sediments. Diatoms were dominant in ice and under-ice water. In the absence of snow, primary production at the end of ice cover period may reach 1 g cal/(m² day). With snow cover present, this index was markedly reduced.     

Shade-adapted Algae beneath Ice and Snow in the Northern Bothnian Sea

The winter-spring phytoplankton flora in the coastal waters of the northern Bothnian Sea consists mainly of marine cold water species even adapted to low light. These dinoflagellates and diatoms seem to be selectively promoted by the seven months of low light and water temperature. They are able to grow during this period of subsaturated light and have their abundance maxima around ice-break. Solar radiation measured under ice and snow cover was in the same order of magnitude as the compensation points of some of the species. In summer, these algae species vanish from the water, and surely survive in the form of inactive life stages.     

Plankton ecology in an ice-covered bay of Lake Michigan: utilization of a winter phytoplankton bloom by reproducing copepods

Plankton ecology was examined during the 1986 winter in Grand Traverse Bay, a 190 m deep, fjordlike bay on Lake Michigan. Before ice cover, algal concentration was low and uniformly distributed with depth, as it is in open Lake Michigan. During ice cover (February and March), a bloom of a typical winter-spring phytoplankton community developed in the upper 40 m, resulting in a 4 to 7-fold increase in feeding rate of adult Diaptomus spp. High algal concentration and zooplankton feeding persisted after ice melt (April). During and after ice cover, lipid concentrations of Diaptomus dropped rapidly from 34% of dry weight to 17 % because of egg production. High incident photosynthetically active radiation (PAR), high (45–50%) PAR transmittance of the ice due to little snow on the ice, and water column stability were probably responsible for the bloom. High ice transparency may be a common feature of large lakes and bays, where strong winds blow snow cover off the ice, or at low latitudes where snowmelt due to occasional rains and warm temperature is common. Winter reproducing calanoid copepods use these blooms to increase their reproductive output.     

SHADE ADAPTED BENTHIC DIATOMS BENEATH ANTARCTIC SEA ICE1

A dense community of shade adapted microalgae dominated by the diatom Trachyneis aspera is associated with a siliceous sponge spicule mat in McMurdo Sound, Antarctica. Diatoms at a depth of 20 to 30 m were found attached to spicule surfaces and in the interstitial water between spicules. Ambient irradiance was less than 0.6 μE · m^?2· s^?1 due to light attenuation by surface snow, sea ice, ice algae, and the water column. Photosynthesis-irradiance relationships determined by the uptake of NaH¹⁴CO₃ revealed that benthic diatoms beneath annual sea ice were light-saturated at only 11 μE·m^?2·s^?1, putting them among the most shade adapted microalgae reported. Unlike most shade adapted microalgae, however, they were not photoinhibited even at irradiances of 300 μE·m^?2·s^?1. Although in situ primary production by benthic diatoms was low, it may provide a source of fixed carbon to the abundant benthic invertebrates when phytoplankton or ice algal carbon is unavailable.     

Vertical and seasonal variation of phytoplankton photosynthesis in a brown-water lake with winter ice cover

SUMMARY. Unlike previously studied lakes with prolonged winter ice and snow cover, Lake Paajarvi, southern Finland, has a high humus content and consequently differs in both the quantity and quality of light penetration into its waters. Moreover, the range of temperature fluctuation and the degree of development of thermal stratification are greater in Paajarvi, and this increased environmental heterogeneity apparently stimulates diversity in the phytoplankton community, especially in the seasonal succession of species. Differences in the photosynthetic capacity of algae from different depths in the water column were not great. This is attributed to the extremely shallow euphotic zone, algae circulating freely through the steep light gradient and sedimenting rapidly once they pass through the thermocline into the hypolimnion. It is suggested that 'adaptation' of phytoplankton to the great seasonal changes in irradiance is achieved largely by successive growths of different species in the community, and that the adaptations and vertical migrations by individual algal species, which have been reported from polar and high alpine lakes, may be of secondary importance in Pääjärvi. The species successions in Pääjärvi produce changes in the pigment content of algae similar to those reported from polar and high alpine lakes, confirming that change in pigmentation is an important mechanism in light adaptation, whether at community or individual level. Algal pigment content was particularly high at the end of the long period of winter ice cover, indicating a degree of adaptation to the prolonged low-light conditions, which produced the extremely high photosynthetic capacities measured at this time. However, phytoplankton production at any irradiance was primarily determined by biomass.     

Winter phytoplankton communities in different depths of three mesotrophic lakes (Łęczna-Włodawa Lakeland, Eastern Poland)

Phytoplankton samples were collected from three mesotrophic lakes: Piaseczno, Rogóźno and Krasne during winter seasons (from January to March). The samples were analyzed for species analysis and abundance of planktonic algae in relation to different depths of water column (0–7 m). Selected water physical-chemical parameters were also measured. Abundance of phytoplankton depended strongly on the thickness of snow and ice cover or mixing conditions. The maximal phytoplankton total number reached about 5 × 10⁶ ind. L⁻¹ beneath the clear ice in the Krasne Lake, minimal numbers were recorded under the thick snow and ice layers in the Piaseczno Lake (2 × 10³ ind. L⁻¹). The winter phytoplankton communities were dominated by flagellates principally cryptomonads (Cryptomonas spp. Rhodomonas minuta), euglenophytes (Trachelomonas volvocina, T. volvocinopsis), dinoflagellates (Peridinium bipes, Gymnodinium helveticum) and chrysophytes (Mallomonas elongata, M. akrokomos, Dinobryon sociale) or non-motile small species of blue-green algae (e.g. Rhabdoderma lineare, Limnothrix redekei), diatoms (Stephanodiscus spp., Asterionella formosa), and green algae (e.g. Scenedesmus spp., Monoraphidium spp.). Phytoplankton abundance and structure showed differentiation during the winter season and along the water column as well.     

Changes in Phytoplankton Biomass and Nutrient Quantities in Sea Ice as Responses to Light/Dark Manipulations during different Phases of the Baltic Winter 2003

The response of Baltic Sea ice communities to changing light climate was studied in three subsequent 3 week in situ experiments on the SW coast of Finland. The investigation covered three different winter periods, short day with low solar angles leading to limited light in the ice, late winter with deep snow cover and early spring with melting snow and increasing light availability. The experimental setup consisted of transparent (no snow) and completely darkened (heavy snow cover) plexiglass tubes in which the ice cores were incubated in situ from 1 to 2 weeks. Changes in the concentrations of inorganic nutrients (NO₃⁻-–N, PO₄³⁻-–P, SiO₄⁻-–Si) and chlorophyll-a concentration in the phytoplankton community composition were recorded as responses to different light manipulations. Changes in inner ice light intensity in untreated ice as well as the temperature both in air and ice were recorded over the entire study period. Increased irradiance in late winter/early spring and during meltdown affected the chlorophyll-a amount in the sea ice. During these periods the phytoplankton community in the top layers decreased possibly as a consequence of photo-acclimation. Closer to the bottom of the ice, however, the increased inner ice light intensity induced algal growth. Complete exclusion of light stopped the algal growth in the whole ice column. Darkening the ice cores also slowed down the ice melting opposite to accelerated melting caused by increased light. The significant differences found in nutrient concentrations between the light and dark treatments were mostly explicable by changes in algal biomass. No obvious changes were observed in the phytoplankton community composition due to light manipulation, diatoms and heterotrophic flagellates dominating throughout the study period.     

Annual cycle of phytoplankton in Ace Lake,an ice covered,saline meromictic lake

The annual cycle of phytoplankton in saline, meromictic Ace Lake (68°2S.4S, 78°11.1E) in the Vestfold Hills, Antarctica, was studied from January, 1979 to January 1980. Ace Lake has permanent gradients of temperature, salinity, dissolved oxygen, and hydrogen sulphide, and is ice covered with up to 2 m of ice for 10–12 months each year. The phytoplankton community had low diversity, consisting of only four species, all flagellates — a prasinophyte Pyramimonas gelidicola McFadden et al., a cryptophyte of the genus Cryptomonas; an unidentified colourless microflagellate, and an unarmoured dinoflagellate. These were restricted to the oxic zone of the lake from the surface to 10 m.The phytoplankton had a cycle of seven months of active growth over spring and summer. Low numbers of cells survived in the water column over winter. Spring growth was initiated below the ice by increased light penetration through the ice into the lake, enhanced at the time by the removal of surface snow which accumulated on the ice over winter. Peak phytoplankton biomass production was by the shade adapted P. gelidicola and occurred at the interface of the oxic and anoxic zones where substantial available nitrogen as ammonia is found.The three dominant phytoplankton species displayed distinct vertical stratification over the oxic zone. This stratification was not static and developed over spring as the flagellates migrated to preferred light climate zones. Mean cell volume of two of the flagellates varied significantly over the year. Minimum volumes were recorded in winter and volume increased progressively over spring to reach maximum mean cell volume in summer. Mean cell volume was positively correlated with light intensity (maximum ambient PAR at the respective depth for date of sample). Low cell volume in winter may be related to winter utilization of carbohydrate reserves by slow respiration, and may represent a survival mechanism.     

THE EFFECTS OF ULTRAVIOLET‐B RADIATION ON ANTARCTIC SEA‐ICE ALGAE1

The impacts of ultraviolet‐B radiation (UVB) on polar sea‐ice algal communities have not yet been demonstrated. We assess the impacts of UV on these communities using both laboratory experiments on algal isolates and by modification of the in situ spectral distribution of the under‐ice irradiance. In the latter experiment, filters were attached to the upper surface of the ice so that the algae were exposed in situ to treatments of ambient levels of PAR and UV radiation, ambient radiation minus UVB, and ambient radiation minus all UV. After 16 d, significant increases in chl a and cell numbers were recorded for all treatments, but there were no significant differences among the different treatments. Bottom‐ice algae exposed in vitro were considerably less tolerant to UVB than those in situ, but this tolerance improved when algae were retained within a solid block of ice. In addition, algae extracted from brine channels in the upper meter of sea ice and exposed to PAR and UVB in the laboratory were much more tolerant of high UVB doses than were any bottom‐ice isolates. This finding indicates that brine algae may be better adapted to high PAR and UVB than are bottom‐ice algae. The data indicate that the impact of increased levels of UVB resulting from springtime ozone depletion on Antarctic bottom‐ice communities is likely to be minimal. These algae are likely protected by strong UVB attenuation by the overlying ice and snow, by other inorganic and organic substances in the ice matrix, and by algal cells closer to the surface.     

Ecology under lake ice

Winter conditions are rapidly changing in temperate ecosystems, particularly for those that experience periods of snow and ice cover. Relatively little is known of winter ecology in these systems, due to a historical research focus on summer ‘growing seasons’. We executed the first global quantitative synthesis on under‐ice lake ecology, including 36 abiotic and biotic variables from 42 research groups and 101 lakes, examining seasonal differences and connections as well as how seasonal differences vary with geophysical factors. Plankton were more abundant under ice than expected; mean winter values were 43.2% of summer values for chlorophyll a, 15.8% of summer phytoplankton biovolume and 25.3% of summer zooplankton density. Dissolved nitrogen concentrations were typically higher during winter, and these differences were exaggerated in smaller lakes. Lake size also influenced winter‐summer patterns for dissolved organic carbon (DOC), with higher winter DOC in smaller lakes. At coarse levels of taxonomic aggregation, phytoplankton and zooplankton community composition showed few systematic differences between seasons, although literature suggests that seasonal differences are frequently lake‐specific, species‐specific, or occur at the level of functional group. Within the subset of lakes that had longer time series, winter influenced the subsequent summer for some nutrient variables and zooplankton biomass.     

Winter aggregations of marine mammals and birds in the north-eastern Weddell Sea pack ice

Summary A seabird and mammal census was carried out in the north-eastern Weddell Sea during the austral winter of 1986. The German research icebreaker Polarstern operated in heavy pack ice along the Greenwich Meridian between the northern sea ice boundary and the Antarctic coast. Crabeater seals (Lobodon carcinophagus), minke whales (Balaenoptera acutorostrata), Adélie penguins (Pygoscelis adeliae), Antarctic petrels (Thalassoica antarctica) and snow petrels (Pagodroma nivea) were found to be more abundant in the vicinity of the submarine Maud Rise, about 700 km north of the continental margin, than in other areas of substantial ice cover traversed during that cruise. The aggregations of birds and mammals are expected to reflect aggregations of their principal food, krill (Euphausia superba) wintering underneath the ice cover. The distribution pattern of krill predators coincides with the course of a warm water belt upwelling near Maud Rise. This upwelling could induce local ice melting which in turn may result in an increased release of sea ice algae.     

Phytoplankton of the Barents Sea - the end of a growth season

 Few phytoplankton investigations have been carried out at the end of the growth season, particularly in the Arctic. In the present study, we monitored the phytoplankton distribution in relation to environmental conditions in the Barents Sea in September 1988 and October 1987. An ice-edge bloom was found in September at 80° N in a stratified meltwater layer, lasting until new ice formation and southward advection of the ice cover commenced in the middle of the month. Phytoplankton populations in the marginal ice zone at this time were not nutrient limited, but biomass was probably reduced due to grazing by small copepods. Lower chl/C and chl/N ratios in the phytoplankton above the pycnocline than below in September indicated light-adapted populations. In October the particulate matter was rich in carbon, but had low chlorophyll content, indicating high levels of detritus. The hydrographic conditions in October differed greatly from those observed in September. The combination of freezing and mixing resulted in higher salinity and nutrient concentrations, and caused a homogeneous distribution, as well as reduction, of the phytoplankton stocks in the upper water column. During late October, low incoming radiation, combined with deep vertical mixing, resulted in light-limiting conditions for the algae, eventually stopping photosynthesis and terminating the growth season in the northern Barents Sea. Received: 1 March 1996/Accepted: 19 May 1996     

Effects of cooling water discharge on the structure and dynamics of epilithic algal communities in the northern Baltic

The Forsmark Biotest Basin is a shallow coastal ecosystem that receives brackish cooling-water discharge from a nuclear power plant. The effects of the discharge on epilithic algal communities were investigated by analysing samples taken every third week throughout one year at 11 sites differentially affected by temperature and/or flow rate enhancement. Community variation was summarized in a canonical correspondence analysis (CCA) of species abundances as a function of site and date. The temperature increase favoured blue-green algae at the expense of red and brown algae. Blue-green algae were however abundant in summer in stagnant water, whether heated or not, and some red and brown algae became abundant in winter in heated sites with flowing water. Green algae and diatoms increased in biomass in the heated sites, but not in relative cover-abundance. The absence of ice and snow cover at sites with heated and/or flowing water caused autumn species to persist into winter, because of the higher light intensity (compared with natural conditions) and the absence of the mechanical abrasion by ice. The thermal discharge lowered species diversity (Shannon-Weaver index) both in summer and winter at sites with flowing water, but not at sites with quiescent or stagnant water. CCA showed alternate periods of stability and rapid change within the seasonal cycle. Individual species were placed according to their optimum; red and brown algae in winter/spring, green algae in spring/summer, blue-green algae in summer, and diatoms at various times. Exceptions to this pattern were species endo- or epiphytic on species of a different group. Analysis of the effects of temperature, flow rate and ice cover on the seasonal pattern of particular species showed that different species respond in individualistic ways to different combinations of these environmental variables.     

Dynamics of ice algae and phytoplankton in Frobisher Bay

Summary Vertical and seasonal variations of ice algae and phytoplankton were studied in relation to their physico-chemical environments in Frobisher Bay from 1979 to 1986. The biomass, estimated by both chlorophyll a concentrations and cell counts, was greater in the ice algae than in the phytoplankton in the underlying sea-water during winter and spring. Algal distribution in the sea ice varied vertically and seasonally, while in the underlying water column the phytoplankton distribution was much less variable. The ice algal bloom occurred at the bottom of the ice, particularly in the lower 5 cm during late spring, while the phytoplankton bloom took place at depths between 1 and 10 m during early summer after the ice bloom was over. The community structure of the ice algae changed from pennate to centric diatoms as the ice melted. The centrics dominated through the fall, and then decreased as the pennates increased in dominance when the ice formed again in winter. Species diversity and number were greater in the sea ice than in the seawater, but they were similar vertically within each habitat. The evenness of the species distribution did not vary with ice thickness or water depth. Species composition, abundance and dominance of ice algae and phytoplankton continually change both vertically and seasonally. The differential abilities of the species to attain maximal growth rates under various environmental conditions may result in species succession. Evidence is given for the major role of environmental factors regulating the dynamics of ice algae and phytoplankton.     

Effects of snow removal and algal photoacclimation on growth and export of ice algae

Net growth of ice algae in response to changes in overlying snow cover was studied after manipulating snow thickness on land-fast, Arctic sea ice. Parallel laboratory experiments measured the effect of changing irradiance on growth rate of the ice diatom, Nitzschia frigida. After complete removal of thick snow (≥9 cm), in situ ice algae biomass declined (over 7–12 days), while removal of thin snow layers (4–5 cm), or partial snow removal, increased net algal growth. Ice bottom ablation sometimes followed snow removal, but did not always result in net loss of algae. Similarly, in laboratory experiments, small increases in irradiance increased algal growth rate, while greater light shifts suppressed growth for 3–6 days. However, N. frigida could acclimate to relatively high irradiance (110 μmol photons m² s⁻¹). The results suggest that algal loss following removal of a thick snow layer was due to the combination of photoinhibition and bottom ablation. The smaller relative increase in irradiance after removal of thin or partial snow layers allowed algae to maintain high specific-growth rates that compensated for loss from physical mechanisms. Thus, the response of ice algae to snow loss depends both on the amount of change in snow depth and algal photophysiology. The complex response of ice algae growth and export loss to frequently changing snow fields may contribute to horizontal and temporal patchiness of ecologically and biogeochemically important variables in sea ice and should be considered in predictions of how climate change will affect Arctic marine ecosystems.     

Cumulative solar irradiance and potential large-scale sea ice algae distribution off East Antarctica (30°E–150°E)

We present a computational model of the large-scale cumulative light exposure of sea ice in the Southern Ocean off East Antarctica (30°E–150°E). The model uses remotely sensed or modelled sea ice concentration, snow depth over sea ice, and solar irradiance data, and tracks sea ice motion over the season of interest in order to calculate the cumulative exposure of the ice field to photosynthetically active radiation (PAR). Light is the limiting factor to sea ice algal growth over winter and early spring, and so the results have implications for the estimation of algal biomass in East Antarctica. The model results indicate that highly light-exposed ice is restricted to within a few degrees of the coast in the eastern part of the study region, but extends much further north in the 30°E–100°E sector. The relative influences of sea ice motion, solar flux, and snow depth variations on interannual variations in model predictions were evaluated. The model estimates of cumulative PAR were found to correlate with satellite estimates of subsequent open-water chlorophyll-a concentration, consistent with the notion that sea ice algae can provide inocula for phytoplankton blooms.     

Experimental Evidence on Nutrient and Substrate Limitation of Baltic Sea sea-ice Algae and Bacteria

The effect of nutrient limitation on Baltic Sea ice algae, and substrate and nutrient limitation on ice bacteria, was studied in a series of in situ -experiments conducted during the winter of 2002 in northern Baltic Sea. Community level changes in algal biomass (chlorophyll a) and productivity, and bacterial thymidine and leucine incorporation were followed for one week after the addition of nutrient and/or organic carbon rich filtered seawater to the experimental units. The results showed the major contribution of snow cover to the algal responses during the beginning of the ice-covered season. Algal communities were able to grow even in January if no snow was present. Nutrient addition did occasionally have an effect on algal biomass and productivity in the ice. Surprisingly, seeding effect from the ice to the underlying water was negatively affected by the nutrient availability in March. Bacterial limitation varied between nutrient (phosphorus) and substrate limitations. The results showed, that limitation in both algal and bacterial communities changed periodically in the northern Baltic Sea ice.     

PRESENCE OF ALGAE IN FRESHWATER ICE COVER OF FLUVIAL LAC SAINT‐PIERRE (ST. LAWRENCE RIVER,CANADA)1

Winter ice cover is a fundamental feature of north temperate aquatic systems and is associated with the least productive months of the year. Here we describe a previously unknown freshwater habitat for algal and microbial communities in the ice cover of the freshwater St. Lawrence River, Quebec, Canada. Sampling performed during winter 2005 revealed the presence of viable algal cells, such as Aulacoseira islandica (O. Müll.) Simonsen (Bacillariophyceae), and microbial assemblage growing in the ice and at the ice–water interface. Vertical channels (1–5 mm wide) containing algae were also observed. Concentrations of chl a ranged between 0.5 and 169 μg · L^?1 of melted ice, with maximal concentrations found in the lower part of the ice cores. These algae have the potential to survive when ice breakup occurs and reproduce rapidly in spring/summer conditions. Freshwater ice algae can thus contribute to in situ primary production, biodiversity, and annual carbon budget in various habitats of riverine communities.     

Light‐driven tipping points in polar ecosystems

Some ecosystems can undergo abrupt transformation in response to relatively small environmental change. Identifying imminent ‘tipping points’ is crucial for biodiversity conservation, particularly in the face of climate change. Here, we describe a tipping point mechanism likely to induce widespread regime shifts in polar ecosystems. Seasonal snow and ice‐cover periodically block sunlight reaching polar ecosystems, but the effect of this on annual light depends critically on the timing of cover within the annual solar cycle. At high latitudes, sunlight is strongly seasonal, and ice‐free days around the summer solstice receive orders of magnitude more light than those in winter. Early melt that brings the date of ice‐loss closer to midsummer will cause an exponential increase in the amount of sunlight reaching some ecosystems per year. This is likely to drive ecological tipping points in which primary producers (plants and algae) flourish and out‐compete dark‐adapted communities. We demonstrate this principle on Antarctic shallow seabed ecosystems, which our data suggest are sensitive to small changes in the timing of sea‐ice loss. Algae respond to light thresholds that are easily exceeded by a slight reduction in sea‐ice duration. Earlier sea‐ice loss is likely to cause extensive regime shifts in which endemic shallow‐water invertebrate communities are replaced by algae, reducing coastal biodiversity and fundamentally changing ecosystem functioning. Modeling shows that recent changes in ice and snow cover have already transformed annual light budgets in large areas of the Arctic and Antarctic, and both aquatic and terrestrial ecosystems are likely to experience further significant change in light. The interaction between ice‐loss and solar irradiance renders polar ecosystems acutely vulnerable to abrupt ecosystem change, as light‐driven tipping points are readily breached by relatively slight shifts in the timing of snow and ice‐loss.