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Ice algae
Ice algae
Ice algae
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Ice algae are any of the various types of algal communities found in annual and multi-year sea, and terrestrial lake ice or glacier ice.

On sea ice in the polar oceans, ice algae communities play an important role in primary production.[1] The timing of blooms of the algae is especially important for supporting higher trophic levels at times of the year when light is low and ice cover still exists. Sea ice algal communities are mostly concentrated in the bottom layer of the ice, but can also occur in brine channels within the ice, in melt ponds, and on the surface.

Because terrestrial ice algae occur in freshwater systems, the species composition differs greatly from that of sea ice algae. In particular, terrestrial glacier ice algae communities are significant in that they change the color of glaciers and ice sheets, impacting the reflectivity of the ice itself.

Sea ice algae

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Antarctic diatom algae covering the underwater surface of broken sea ice in the Ross Sea.

Adapting to the sea ice environment

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Microbial life in sea ice is extremely diverse,[2][3][4] and includes abundant algae, bacteria and protozoa.[5][6] Algae in particular dominate the sympagic environment, with estimates of more than 1000 unicellular eukaryotes found to associate with sea ice in the Arctic.[7][4][3][2] Species composition and diversity vary based on location, ice type, and irradiance. In general, pennate diatoms such as Nitzschia frigida[8][9] (in the Arctic)[10] and Fragilariopsis cylindrus (in the Antarctic)[11] are abundant. Melosira arctica, which forms up to meter-long filaments attached to the bottom of the ice, are also widespread in the Arctic and are an important food source for marine species.[11]

While sea ice algae communities are found throughout the column of sea ice, abundance and community composition depends on the time of year.[12] There are many microhabitats available to algae on and within sea ice, and different algal groups have different preferences. For example, in late winter/early spring, motile diatoms like N. frigida have been found to dominate the uppermost layers of the ice, as far as briny channels reach, and their abundance is greater in multi-year ice (MYI) than in first year ice (FYI). Additionally, dinoflagellates have also been found to dominant in the early austral spring in Antarctic sea ice.[5]

Sea ice algal communities can also thrive at the surface of the ice, in surface melt ponds, and in layers where rafting has occurred. In melt ponds, dominant algal types can vary with pond salinity, with higher concentrations of diatoms being found in melt ponds with higher salinity.[13] Because of their adaption to low light conditions, the presence of ice algae (in particular, vertical position in the ice pack) is primarily limited by nutrient availability. The highest concentrations are found at the base of the ice because the porosity of that ice enables nutrient infiltration from seawater.[14]

To survive in the harsh sea ice environment, organisms must be able to endure extreme variations in salinity, temperature, and solar radiation. Algae living in brine channels can secrete osmolytes, such as dimethylsulfoniopropionate (DMSP), which allows them to survive the high salinities in the channels after ice formation in the winter, as well as low salinities when the relatively fresh meltwater flushes the channels in the spring and summer. Some sea ice algae species secrete ice-binding proteins (IBP) as a gelatinous extracellular polymeric substance (EPS) to protect cell membranes from damage from ice crystal growth and freeze thaw cycles.[15] EPS alters the microstructure of the ice and creates further habitat for future blooms. Ice algae survive in environments with little to no light for several months of the year, such as within ice brine pockets. Such algae have specialized adaptations to be able to maintain growth and reproduction during periods of darkness. Some sea ice diatoms have been found to utilize mixotrophy when light levels are low. For example, some Antarctic diatoms downregulate glycolysis in environments with low to no irradiance, while upregulating other mitochondrial metabolic pathways, including the Entner−Doudoroff pathway which provides the TCA cycle (an important component in cellular respiration) with pyruvate when pyruvate cannot be obtained via photosynthesis.[16] Surface-dwelling algae produce special pigments to prevent damage from harsh ultraviolet radiation. Higher concentrations of xanthophyll pigments act as a sunscreen that protects ice algae from photodamage when they are exposed to damaging levels of ultraviolet radiation upon transition from ice to the water column during the spring.[3] Algae under thick ice have been reported to show some of the most extreme low light adaptations ever observed. They are able to perform photosynthesis in an environment with just 0.02% of the light at the surface.[17] Extreme efficiency in light utilization allows sea ice algae to build up biomass rapidly when light conditions improve at the onset of spring.[18]

Role in ecosystem

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Sea ice algae play a critical role in primary production and serve as part of the base of the polar food web by converting carbon dioxide and inorganic nutrients to oxygen and organic matter through photosynthesis in the upper ocean of both the Arctic and Antarctic. Within the Arctic, estimates of the contribution of sea ice algae to total primary production ranges from 3-25%, up to 50-57% in high Arctic regions.[19][20] Sea ice algae accumulate biomass rapidly, often at the base of sea ice, and grow to form algal mats that are consumed by amphipods such as krill and copepods. Ultimately, these organisms are eaten by fish, whales, penguins, and dolphins.[18] When sea ice algal communities detach from the sea ice they are consumed by pelagic grazers, such as zooplankton, as they sink through the water column and by benthic invertebrates as they settle on the seafloor.[3] Sea ice algae as food are rich in polyunsaturated and other essential fatty acids, and are the exclusive producer of certain essential omega-3 fatty acids that are important for copepod egg production, egg hatching, and zooplankton growth and function.[3][21]

The undersurface of the pack ice in Antarctica colored green - Antarctic krill scraping off the ice algae

Temporal variation

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The timing of sea ice algae blooms has a significant impact on the entire ecosystem. Initiation of the bloom is primarily controlled by the return of the sun in the spring (i.e. the solar angle). Because of this, ice algae blooms usually occurs before the blooms of pelagic phytoplankton, which require higher light levels and warmer water.[21] Early in the season, prior to the ice melt, sea ice algae constitute an important food source for higher trophic levels.[21] However, the total percentage that sea ice algae contribute to the primary production of a given ecosystem depends strongly on the extent of ice cover. The thickness of snow on the sea ice also affects the timing and size of the ice algae bloom by altering light transmission.[22] This sensitivity to ice and snow cover has the potential to cause a mismatch between predators and their food-source, sea ice algae, within the ecosystem. This so called match/mismatch has been applied to a variety of systems.[23] Examples have been seen in the relationship between zooplankton species, which rely on sea ice algae and phytoplankton for food, and juvenile walleye pollock in the Bering Sea.[24]

Bloom initialization

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There are several ways in which sea ice algal blooms are thought to start their annual cycle, and hypotheses about these vary depending on water column depth, sea ice age, and taxonomic group. Where sea ice overlays deep ocean, it is proposed that cells trapped in multiyear ice brine pockets are reconnected to the water column below and quickly colonize nearby ice of all ages. This is known as the multiyear sea ice repository hypothesis.[12] This seeding source has been demonstrated in diatoms, which dominate sympagic blooms. Other groups, such as the dinoflagellates, which also bloom in the spring/summer, have been shown to maintain low cell numbers in the water column itself, and do not primarily overwinter within the ice.[25] Where sea ice covers ocean that is somewhat shallower, resuspension of cells from the sediment may occur.[26]

Implications of climate change

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Climate change and warming of Arctic and Antarctic regions have the potential to greatly alter ecosystem functioning. Decreasing ice cover in polar regions is expected to lessen the relative proportion of sea ice algae production to measures of annual primary production.[27][28] Thinning ice allows for greater production early in the season but early ice melting shortens the overall growing season of the sea ice algae. This melting also contributes to stratification of the water column that alters the availability of nutrients for algae growth by decreasing the depth of the surface mixed layer and inhibiting the upwelling of nutrients from deep waters. This is expected to cause an overall shift towards pelagic phytoplankton production.[28] Changes in multiyear ice volume[29] will also have an impact on ecosystem function in terms of bloom seeding source adjustment. Reduction in MYI, a temporal refugia for diatoms in particular, will likely alter sympagic community composition, resulting in bloom initialization that derives from species that overwinter in the water column or sediments instead.[25]

Because sea ice algae are often the base of the food web, these alterations have implications for species of higher trophic levels.[19] The reproduction and migration cycles of many polar primary consumers are timed with the bloom of sea ice algae, meaning that a change in the timing or location of primary production could shift the distribution of prey populations necessary for significant keystone species. Production timing may also be altered by the melting through of surface melt ponds to the seawater below, which can alter sea ice algal habitat late in the growing season in such a way as to impact grazing communities as they approach winter.[30]

The production of DMSP by sea ice algae also plays an important role in the carbon cycle. DMSP is oxidized by other plankton to dimethylsulfide (DMS), a compound which is linked to cloud formation. Because clouds impact precipitation and the amount of solar radiation reflected back to space (albedo), this process could create a positive feedback loop.[31] Cloud cover would increase the insolation reflected back to space by the atmosphere, potentially helping to cool the planet and support more polar habitats for sea ice algae. As of 1987, research has suggested that a doubling of cloud-condensation nuclei, of which DMS is one type, would be required to counteract warming due to increased atmospheric CO2 concentrations.[32]

Sea ice algae as a tracer for paleoclimate

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Sea ice plays a major role in the global climate.[33] Satellite observations of sea ice extent date back only until the late 1970s, and longer term observational records are sporadic and of uncertain reliability.[34] While terrestrial ice paleoclimatology can be measured directly through ice cores, historical models of sea ice must rely on proxies.

Organisms dwelling on the sea ice eventually detach from the ice and fall through the water column, particularly when the sea ice melts. A portion of the material that reaches the seafloor is buried before it is consumed and is thus preserved in the sedimentary record.

There are a number of organisms whose value as proxies for the presence of sea ice has been investigated, including particular species of diatoms, dinoflagellate cysts, ostracods, and foraminifers. Variation in carbon and oxygen isotopes in a sediment core can also be used to make inferences about sea ice extent. Each proxy has advantages and disadvantages; for example, some diatom species that are unique to sea ice are very abundant in the sediment record, however, preservation efficiency can vary.[35]

Terrestrial snow and ice algae

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Lake snow and ice algae Algae can grow within and attached to lake ice as well, especially below clear, black ice.[36] Within the ice, algae often grows in water-filled air pockets found in the slush layer formed between the ice and snow interface.[37] For instance, the diatom species Aulacoseira baicalensis endemic to Lake Baikal can reproduce intensively in water-filled pockets within the ice as well as attached to the ice sheet.[36] Alpine freshwater ice and snow which can last over half a year has been found to support an overall higher microbial biomass and algal activity than the lake water itself as well as specific predatory species of ciliates only found in the slush layer of the ice and snow interface.[38] Algae living on the snowpack of ice-covered lakes may be especially rich in essential polyunsaturated fatty acids.[39]

Snow and glacier Ice algae Algae also thrive on snow fields, glaciers and ice sheets. The species found in these habitats are distinct from those associated with sea ice because the system is freshwater and the algae are pigmented. Even within these habitats, there is a wide diversity of habitat types and algal assemblages that colonize snow and ice surfaces during melt. For example, cryosestic communities are specifically found on the surface of glaciers where the snow periodically melts during the day.[40] Research has been done on glaciers and ice sheets across the world and several species have been identified. However, although there seems to be a wide array of species they have not been found in equal amounts. The most abundant species identified on different glaciers are the glacier ice algae Ancylonema nordenskioldii[41][42][43][44] and the snow algae Chlamydomonas nivalis.[44][45][46]


Table 1. Algae Species Composition Across Studies on Glaciers and Ice Sheets

Genus Species Source
Mesotaenium bregrenii [42][44]
Ancylonema nordenskioldii [41][42][44]
Cylindrocystis brebissonii [42][44]
Chlamydomonas nivalis [44][45][46]
Phromidemis priestleyi [44]
Oscillatoriaceae cyanobacterium [44]
Chlorooceae cyanobacterium [44]
Chroococcaceae cyanobacterium [41][45]
Chloroplastida [46]
Chloromonas polyptera [46]
Chlamydomonas alpina [46]
Chlamydomonas tughillensis [46]

Implications for climate change

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The rate of glacier melt depends on the surface albedo. Recent research has shown the growth of snow and glacier ice algae darkens local surface conditions, decreasing the albedo and thus increases the melt rate on these surfaces.[46][45][47] Melting glaciers and ice sheets have been directly linked to increase in sea level rise.[48] The second largest ice sheet is the Greenland Ice Sheet which has been retreating at alarming rates. Sea level rise will lead to an increase in both frequency and intensity of storm events.[48]

On enduring ice sheets and snow pack, terrestrial ice algae often color the ice due to accessory pigments, popularly known as "watermelon snow". The dark pigments within the structure of algae increases sunlight absorption, leading to an increase in the melting rate.[41] Algae blooms have been shown to appear on glaciers and ice sheets once the snow had begun to melt, which occurs when the air temperature is above the freezing point for a few days.[45] The abundance of algae changes with the seasons and also spatially on glaciers. Their abundance is highest during the melting season of glaciers which occurs in the summer months.[41] Climate change is affecting both the start of the melting season and also the length of this period, which will lead to an increase in the amount of algae growth.

Ice–albedo feedback loop (SAF)

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Main article: Ice–albedo feedback

As the ice/snow begins to melt the area the ice covers decreases which means a higher portion of land is exposed. The land underneath the ice has a higher rate of solar absorption due to it being less reflective and darker. Melting snow also has lower albedo than dry snow or ice because of its optical properties, so as snow begins to melt the albedo decreases, which results in more snow melting, and the loop continues. This feedback loop is referred to as the Ice–albedo feedback loop. This can have drastic effects on the amount of snow melting each season. Algae plays a role in this feedback loop by decreasing the level of albedo of the snow/ice. This growth of algae has been studied but its exact effects on decreasing albedo is still unknown.

The Black and Bloom project is conducting research to determine the amount algae are contributing to the darkening of the Greenland Ice Sheet, as well as algae's impact on the melting rates of the ice sheets.[49] It is important to understand the extent to which algae is changing the albedo on glaciers and ice sheets. Once this is known, it should be incorporated into global climate models and then used to predict sea level rise.

References

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Ice algae

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Ice algae, also known as algae, are sympagic photosynthetic microorganisms—primarily eukaryotic algae such as pennate diatoms (e.g., Nitzschia frigida)—that inhabit in polar regions through colonization of inclusions, surface flooding, or ponds. These communities, often dominated by diatoms in bottom ice layers, exhibit patchy distributions with varying over scales of 5 meters or more. Microbial activity peaks in spring as algae rapidly colonize the bottommost centimeters of ice following the return of , adapting to low-light conditions via efficient light acclimation and uptake. Ecologically, ice algae contribute substantially to in ice-covered polar waters, potentially accounting for up to 60% of total output in such environments and extending biological productivity by 1–3 months beyond open-water blooms. They form the base of sympagic food webs, providing an early-season carbon source for grazers like copepods and nematodes, and upon ice melt, seed under-ice and pelagic populations essential for higher trophic levels including , , and marine mammals. Over 1,000 microalgal species have been documented in , underscoring their and role in cycling and stability.

Definition and Classification

General Characteristics

Ice algae comprise diverse communities of primarily unicellular , including diatoms, dinoflagellates, and chlorophytes, that inhabit , snowpacks, and glacial surfaces in polar and alpine environments. These organisms often form chains, filaments, or colonies within ice matrices, such as brine channels in or melt layers on snow, enabling them to exploit microhabitats with liquid water despite subzero ambient temperatures. Visible blooms of ice algae can discolor ice formations, producing hues from brownish-green in to red or black on glaciers due to accessory pigments like or . As autotrophic primary producers, ice algae conduct using filtered through , achieving rates sufficient to support polar food webs despite low levels below 10 μmol photons m⁻² s⁻¹. Their cells typically measure 5–100 μm in , with siliceous frustules in s providing structural integrity in high-salinity brines reaching 100–150 ppt. Ice algae exhibit psychrophilic traits, maintaining metabolic activity at temperatures as low as -15°C, as observed in . Biomass accumulation in ice algae communities can reach 10–50 g C m⁻² in under favorable conditions, underscoring their role in early-season productivity before open-water dominate. These demonstrate resilience to osmotic stress and , with cellular mechanisms including proteins and compatible solutes that prevent formation within protoplasts.

Major Taxonomic Groups

Ice algae communities are dominated by a few key taxonomic groups adapted to extreme cold, with diatoms (Bacillariophyceae) comprising the primary constituents in habitats, often accounting for the majority of eukaryotic in bottom assemblages. Pennate diatoms, including genera such as (e.g., N. frigida), , and Fragilariopsis, frequently form dense mats due to their ability to anchor within channels and utilize limited . Centric diatoms like Chaetoceros and Thalassiosira also contribute significantly, particularly in early colonization phases, with species such as C. gelidus and T. reported in ice samples from 2006–2007 studies. Dinoflagellates () and flagellates, including prymnesiophytes and prasinophytes, represent secondary groups in , typically comprising smaller fractions of the community but playing roles in interior and surface layers where aids dispersal. These taxa, such as Polarella glacialis among dinoflagellates, exhibit psychrophilic traits enabling survival in low-salinity pockets. In terrestrial snow and glacier systems, dominate, with (e.g., ) and Zygnematales (e.g., Mesotaenium berggrenii, Ancylonema nordenskioeldii) being the most prevalent orders, responsible for characteristic red, green, or black pigmentation in melt layers. These , comprising over 80% of documented snow algal , produce astaxanthin-like pigments for UV protection and light harvesting under cover. Minor contributions come from euglenoids, , chrysophytes, and dinoflagellates, though these are less abundant and habitat-specific.

Habitats and Global Distribution

Sea Ice Ecosystems

Sea ice ecosystems host diverse microbial communities dominated by ice algae, which colonize brine channels, platelet layers, and the undersurface of ice floes in both the Arctic and Antarctic. These habitats form during sea ice formation, trapping microalgae within the ice matrix or allowing attachment to skeletal layers, with distributions varying by ice type—landfast ice supporting higher biomass accumulation compared to drifting pack ice due to stability and nutrient access. In the Arctic, ice algal biomass peaks in spring, reaching 1 to 100 mg chlorophyll a per square meter during summer under suitable light conditions post-snowmelt. Antarctic communities, often dominated by diatoms in platelet ice under fast ice, exhibit spatial heterogeneity, with chlorophyll a concentrations mapped at millimeter scales revealing patchy distributions influenced by brine volume and salinity gradients. Primary production by ice algae constitutes a significant portion of polar marine productivity, with estimates ranging from 4% to 20% of annual totals in ice-covered areas, driven by seasonal light penetration and nutrient from underlying waters. In the sea-ice zone, ice algae account for 12% to 50% of local despite comprising only about 1% ocean-wide, as evidenced by modeling of gross in landfast ice. These blooms, initiating in spring as cover thins, sustain elevated rates—up to 74% of under-ice pelagic production in some locales—before release during melt supports under-ice and marginal blooms. Year-round carbon signatures from ice algae appear in 96% of sampled organisms, indicating persistent trophic transfer. Within polar food webs, ice algae serve as foundational producers, channeling energy to , , and higher trophic levels like and marine mammals, with fatty acid profiles in consumers reflecting up to 50% reliance on ice-derived sources. In the central , key species derive substantial carbon from ice algae, bolstering resilience amid variable ice conditions, though dependency varies—supplementary rather than primary for some under-ice amphipods. Aggregates of algae under summer distribute basin-scale, seeding post-melt pelagic production and influencing nutrient cycling through exudates and . Declining ice extent poses risks to this basal support, yet adaptive under-ice communities may partially offset losses by exploiting extended open water.

Terrestrial Snow and Glacier Systems

Ice algae in terrestrial systems primarily colonize snowpacks and supraglacial surfaces in polar, alpine, and high-mountain environments, where they form visible blooms during melt seasons. These , distinct from communities, thrive in oligotrophic, low-temperature habitats with high light exposure but limited liquid water, often exhibiting pigmented cells for protection against UV radiation and . dominate in ephemeral or perennial snowfields, while glacier algae preferentially inhabit bare ice zones on retreating and ice sheets. Snow algae communities, comprising mainly such as Chlamydomonas nivalis, Sanguina spp., and Chloromonas spp., occur in red, green, or orange blooms on snow surfaces across the , , and temperate mountains. These taxa are distributed globally in regions with persistent snow cover below 10°C, including the Harding Icefield in , , the European Alps, and coastal snowfields, where blooms can cover extensive areas visible via . In , snow algae form patches on snowfields with metabolic activity peaking in austral summer, supported by films and nutrient influx from atmospheric deposition. High-altitude sites like the and host similar communities, with species diversity increasing toward lower latitudes due to varied melt dynamics. Glacier algae, often from Zygnematophyceae like Ancylonema nordenskiöldii and Mesotaenium berggrenii, inhabit cryoconite holes, ice lenses, and surface melt layers on glaciers worldwide, contributing to darkening that enhances solar absorption. These organisms are prevalent on the , Alaskan glaciers such as Gulkana, and Antarctic ice shelves, with blooms accelerating in warming conditions since the 1980s. Distribution patterns show higher abundances on lower-elevation, debris-free ice, as observed in central Asian glaciers and the , where algal cells accumulate in filaments adapted to flowing . In the , including and Greenland, glacier algae co-occur with during transitional melt phases, forming hybrid communities influenced by topography and impurity levels. Both and exhibit cosmopolitan yet habitat-specific distributions, with endemism emerging in isolated systems like nunataks, as revealed by genomic comparisons of ancient and modern samples. Abiotic factors such as depth, melt duration, and gradients dictate patchiness, with opportunistic dominating transient and specialists persisting on perennial . Recent observations indicate expanding ranges poleward and upslope in response to climate shifts, though data gaps persist in understudied regions like the mountains.

Biological Adaptations

Physiological and Morphological Features

Ice algae exhibit diverse morphologies tailored to their icy habitats, with sea ice communities predominantly comprising pennate diatoms characterized by elongated, bilaterally symmetric cells with silica frustules featuring a central raphe for on ice surfaces and within channels. These diatoms often form adhesive colonies or chains that anchor to ice platelets, facilitating attachment in the turbulent bottom layers of . In contrast, glacier and snow algae, such as in the genera Mesotaenium and Ancylonema, display more rounded or filamentous forms with thick cell walls and secondary like for UV protection, enabling surface colonization and red pigmentation in melt layers. Physiologically, ice algae demonstrate adaptations to subzero temperatures, including the production of cryoprotectants such as antifreeze proteins and compatible solutes that maintain and prevent damage. In sea ice diatoms, low photoadaptive indices (I_k around 10–20 μmol photons m⁻² s⁻¹) and optimal irradiances reflect efficiency in capturing diffuse under-ice light, with enhanced light harvesting via fucoxanthin-chlorophyll proteins. They also exhibit prolonged dark survival through accumulation and reduced metabolic rates, allowing persistence during winter months. Glacier algae further adapt via intracellular nutrient storage, stockpiling and to sustain blooms amid episodic inputs, and high desiccation tolerance via extracellular . These traits collectively enable high primary productivity despite extreme osmotic stress from salinities exceeding 100 psu and temperatures as low as -1.8°C in or -20°C in snowpacks.

Molecular Mechanisms for Extremophile Survival

Ice algae, as psychrophilic , rely on specialized molecular mechanisms to withstand subzero temperatures, high fluctuations, and limited light in matrices. Central to their are ice-binding proteins (IBPs), which adsorb irreversibly to surfaces, creating thermal hysteresis and inhibiting recrystallization that could otherwise puncture cell membranes. These proteins, often acquired via , feature diverse folds such as polyproline type II helices and are encoded by expanded families in like the alga Chloromonas sp. ICE-L, enabling adhesion to and modulation of to prevent lethal intracellular formation.30845-9) Membrane lipid composition undergoes adaptive remodeling through upregulation of desaturase enzymes, increasing the proportion of unsaturated s to preserve fluidity and functionality at temperatures approaching -1.8°C in brine channels. In Chloromonas sp. ICE-L, genomic expansions in biosynthesis genes facilitate this desaturation, countering the rigidifying effects of cold on bilayers and maintaining proton leak and transport processes essential for . Complementary cryoprotectants, including extracellular polymeric substances (EPS) rich in and glycoproteins, accumulate to osmotically stabilize cells against brine rejection and act as spacers that deform lattices, reducing spicule penetration.30845-9) At the transcriptional level, rapid shifts in response to cold stress optimize , with psychrophilic algae like exhibiting differential regulation of genes involved in , carbon fixation, and stress response pathways, such as those encoding cold-shock proteins and chaperones that refold misfolded enzymes. Expanded gene families in sea ice diatoms address cold-induced mutagenesis from generated during low-light , while across taxa reinforces substitutions in codon-biased genes for ribosomal proteins and transporters, enhancing translational efficiency in hypothermic conditions. These mechanisms collectively enable sustained viability, with proteomic studies confirming elevated abundances of signaling and nutrient-scavenging proteins in glacier-associated streptophyte algae under perennial ice.

Ecological and Biogeochemical Roles

Primary Production and Carbon Cycling


Ice algae, primarily sympagic communities in sea ice, drive significant primary production in polar regions through photosynthesis, fixing atmospheric CO₂ into biomass despite limited light penetration. In the Arctic Ocean, ice algae contributions to total primary production vary regionally, ranging from under 1% in nutrient-rich coastal zones to as high as 60% in the central basin during late summer (August–September). Under landfast sea ice near Barrow, Alaska, bottom ice algae supplied 74% of under-ice pelagic primary production before the onset of open-water phytoplankton blooms in spring. Quantitatively, ice algal primary production in the is estimated at 28–211 Tg C yr⁻¹, substantially lower than phytoplankton's 355–3,671 Tg C yr⁻¹, reflecting their confinement to habitats and shorter productive periods. In the , long-term modeling indicates average algal gross primary production of 15.5 Tg C yr⁻¹ since 1850, with higher rates in landfast ice zones supporting localized blooms. Snow and algae exhibit lower production rates, often dominated by red-pigmented species like , but contribute modestly to surface carbon fixation in terrestrial systems, with limited empirical quantification compared to marine counterparts. In carbon cycling, ice algae enhance export fluxes by forming aggregates that sink upon ice melt or sloughing, transferring fixed carbon to benthic ecosystems and potentially sequestering it in sediments. Ice-covered areas show elevated particle export efficiency and vertical microbial connectivity, with sympagic carbon signatures detected in 96% of year-round sampled Arctic organisms, underscoring sustained trophic transfer. Early ice breakup amplifies this export, increasing particulate organic carbon delivery to the seafloor and influencing deep-ocean carbon storage amid declining sea ice. Sympagic algae thus play a pivotal role in polar carbon budgets, bridging surface production to subsurface reservoirs despite their episodic nature.

Nutrient Dynamics and Trophic Interactions

Ice algae significantly influence dynamics in polar sea ice ecosystems through active uptake and intracellular storage of key s, including , , , and phosphates, primarily from channels and underlying . During spring blooms, these accumulate quotas exceeding ambient levels by factors of up to 10-fold for and , enabling rapid biomass accumulation despite limitation in ice pores. This storage strategy, observed in sea ice diatoms as of 2025, decouples algal growth from short-term external supply fluctuations driven by rejection during freezing or dilution via melting. Biogeochemical cycling is further modulated by physical processes such as turbulent fluxes at the ice-ocean interface and seasonal dynamics, which replenish depleted pools and export upon ice melt. In landfast , concentrations exhibit strong seasonality, with winter accumulation followed by spring depletion and post-bloom remineralization, underscoring the role of ice algae in regional carbon and budgets. Trophic interactions position ice algae as a foundational source for sympagic and pelagic , particularly in winter when open-water production ceases. (Euphausia superba) rely on ice algal carbon for overwintering, with stable analyses indicating that sympagic production constitutes up to 50% of juvenile diets in ice-covered regions. This carbon flux supports higher trophic levels, including amphipods and copepods, which transfer ice-derived biomass to and seabirds, with under-ice rates estimated at 3-5 mg C m⁻² day⁻¹ for key . However, reliance varies temporally and spatially; summer utilization by and amphipods remains low, suggesting ice algae serve more as a supplementary rather than dominant resource during blooms. Emerging evidence of under-ice enhances availability for algae, potentially amplifying trophic transfers by boosting basal production in -poor waters. Physical structures like sea-ice terraces may modulate these interactions by providing refugia that alter predation risks and grazing efficiency for larval . Overall, ice algae mediate a critical link between cycling and dynamics, sustaining polar ecosystems amid seasonal ice variability.

Environmental Interactions

Albedo Effects and Surface Processes

Ice algae reduce the of ice surfaces primarily through cellular pigments such as and , which enhance absorption of visible and near-infrared wavelengths compared to clean or . Clean typically reflects 50–80% of incident shortwave , but algal blooms can lower this by 3.5–43% in dense patches on bare ice, depending on density and composition. This albedo reduction increases net shortwave absorption, accelerating surface melt rates; for example, ice algal communities on bare ice generate additional melt of 0.17–1.7 cm water equivalent per day, contributing to broader mass loss. On ice caps, diverse blooms dominated by taxa like Ancylonema (Zygnematophyceae) decrease broadband reflectance (350–1000 nm), yielding instantaneous of ~16.6 W m⁻² and contributing 0.45–2.36% to total melt, or ~7 million liters of across 2.7 km² areas in a single season. Surface processes are amplified by algal activity, as pigmentation-induced heating forms weathering crusts and thin liquid water layers that trap light and further suppress via altered scattering properties. In , spring melt triggers surface algal proliferation, promoting early ponding that reduces structural integrity through brine channel expansion and top-down , with up to 13% decline in affected regions. These dynamics establish positive feedbacks, where melt exposes substrates for further colonization, exacerbating energy imbalance and decay.

Proxy Uses in Paleoenvironmental Reconstruction

Ice algae, particularly sympagic , serve as valuable proxies for reconstructing past conditions due to their association with ice habitats and the preservation of their biomarkers and frustules in marine sediments. These organisms thrive in the under-ice environment, where penetration and during seasonal melt support blooms, leading to deposition of diagnostic remains that reflect cover duration, extent, and retreat patterns over timescales from the to glacial-interglacial transitions. Quantitative reconstructions often integrate multiple lines of , such as species assemblages and biomarkers, to infer paleoproductivity and dynamics linked to broader forcings like orbital variations or circulation shifts. A primary biomarker proxy is IP25, a C25 highly branched isoprenoid (HBI) synthesized by specific diatoms, including minor taxa like Haslea kijimae and Pleurosigma staurophorum, during under-ice growth seasons. Detected in sediments via gas chromatography-mass spectrometry, IP25 presence indicates recurrent seasonal rather than cover or ice-free conditions, with concentrations correlating to ice algal productivity influenced by factors such as thickness and melt timing. For semi-quantitative estimates of past concentration, the PIP25 index combines IP25 with underlying open-water markers like brassicasterol, where higher index values denote thicker or more persistent ; calibrations against modern observations yield extents accurate to within 15-20% for mid-to-late records in regions like the and Canadian Arctic Archipelago. Recent Bayesian statistical frameworks have refined IP25 interpretations by accounting for proxy uncertainties and spatial variability, enabling direct comparisons with simulations of minima around 6-8 thousand years ago. In the Southern Ocean, analogous proxies include IPSO25, a mono-unsaturated HBI variant produced by sea ice diatoms such as Fragilariopsis curta and F. cylindrus, which signals past sea ice margins. Sediment core analyses from sites like the reveal IPSO25 fluctuations tracking deglacial ice retreat around 18-14 thousand years ago, with elevated levels corresponding to expanded winter sea ice during Marine Isotope Stage 2. Complementary triene HBIs (e.g., 13-methyl-triene) may indicate marginal ice zones, where ice-edge blooms occur, enhancing resolution for transitional environments. valve counts provide additional qualitative proxies, with taxa like Thalassiosira antarctica (resting spores) denoting prolonged sea ice presence, as their abundances in cores from the correlate with historical satellite-derived extents over the past millennium. Multi-proxy approaches, integrating HBIs with fluxes and geochemical signals like biogenic silica, have reconstructed sea ice variability, revealing expansions during cooler intervals like the (circa 1400-1850 CE). Fossil diatom assemblages from laminated sediments or ice-rafted debris further proxy ice algal contributions to paleoenvironments, with shifts in sympagic species ratios indicating changes in stability and nutrient . For instance, increased Nitzschia frigida relative to pelagic forms in northern North Atlantic cores signals enhanced incursions during the stadial (12.9-11.7 thousand years ago). These records link ice algae proxies to atmospheric teleconnections, such as strengthened influencing Southern upwelling, though interpretations require calibration against modern analogs to mitigate taphonomic biases like dissolution in undersaturated waters. Overall, ice algae-derived proxies have illuminated feedbacks in past climates, informing projections of under anthropogenic warming.

Climate Change Implications

Observed Responses to Ice Loss

In the Arctic, empirical observations indicate that declining sea ice thickness and extent have advanced the phenology of ice algal blooms, primarily through increased light penetration to bottom-ice communities. Field and satellite data from regions such as the Barents Sea and East Greenland Sea document melt onset advancing by 25 to 30 days since 1979, enabling earlier initiation of algal growth under thinning ice covers typically 0.5–1 meter thick. This shift compresses the productive window, with ice algae often reaching peak biomass (measured as chlorophyll-a concentrations up to 100–200 mg m⁻²) 2–4 weeks earlier than in prior decades, followed by rapid release into the underlying water column upon melt. In Antarctic coastal fast ice, reduced ice persistence has similarly prompted earlier under-ice algal development, with in situ measurements from Svalbard fjords and East Antarctica showing bottom communities acclimating to higher irradiances (up to 50–100 µmol photons m⁻² s⁻¹) under thinner ice, resulting in elevated photosynthetic rates and biomass accumulation before seasonal breakup. However, widespread sea ice minima since 2016—such as the record low extent of 1.92 million km² in March 2023—have led to shorter ice seasons (by 20–40 days in some sectors), diminishing overall ice algal standing stocks and shifting production toward pelagic diatoms upon release, as evidenced by community restructuring with diatom rebounds in surface waters. These changes alter carbon flux, with earlier export pulses enhancing benthic subsidies but potentially reducing sympagic (ice-associated) primary production by 20–50% in affected areas. Light regime alterations from ice loss further influence algal physiology, with field spectra measurements revealing reduced blue light transmission (400–500 nm) and enhanced red wavelengths under open water, impacting the efficiency of ice-adapted species like Nitzschia frigida and Fragilariopsis cylindrus, which exhibit optimized absorption in ice-filtered conditions. Observations from Arctic pack ice transects post-2010 minima confirm decreased under-ice algal viability due to abrupt exposure, with photosynthetic quantum yields dropping 10–30% in transitioned communities lacking ice matrix support.

Debates on Feedback Strength and Ecosystem Resilience

The strength of feedbacks involving ice algae, particularly through albedo reduction and enhanced melt rates, remains a subject of debate due to regional variability and modeling uncertainties. Ice algae lower surface by darkening and , absorbing more solar radiation and accelerating melt in a loop; on Greenland's west coast, this biological effect accounts for approximately 10% of total melt. However, quantification at pan- or scales is challenging, with ice algal production estimates ranging from 3–73 Tg C yr⁻¹ in the and 3–24 Tg C yr⁻¹ in the , and models showing no significant long-term trend in production from 1980–2010 despite decline, attributed to high interannual variability and shifting bloom . Critics argue that physical factors like thickness distribution may dampen the overall ice- feedback more than biological darkening amplifies it, while others highlight potential underestimation in coupled models that overlook algal nutrient efficiency and resilience to warming-induced scarcity. Debates on ecosystem resilience center on whether the loss of sympagic (ice-associated) production can be offset by increased pelagic blooms in open water, or if disruptions to timing and will reduce stability. Some evidence suggests compensatory mechanisms, with primary potentially rising due to extended ice-free periods, but phenological mismatches—such as earlier ice algal blooms outpacing grazers—could weaken trophic transfers and export of carbon to deeper layers. In the decline projections vary widely (29–90% summer reduction by 2100 under high emissions), complicating resilience assessments, as key species like show dependence on ice algae for , with limited quantitative on responses indicating vulnerability to synergistic stressors like . While multiyear ice loss reduces algal diversity, potentially eroding resilience, certain benthic communities may benefit from increased light penetration, though overall polar s exhibit lower climatic resilience than previously assumed due to rapid cryospheric shifts.

Research History and Advances

Early Discoveries and Field Observations

The presence of microorganisms, including , within was first documented in the early 1840s through examinations of pack ice samples collected during exploratory voyages. Protozoologist Christian Gottfried Ehrenberg described diverse and algal forms embedded in the ice, noting their viability despite subfreezing conditions, based on microscopic analysis of specimens preserved from northern expeditions. These initial reports established as a for photosynthetic life, though Ehrenberg's interpretations emphasized protozoan dominance over algal components. Subsequent field observations in the late 19th and early 20th centuries expanded on these findings during polar expeditions. Norwegian botanist Hans Henrik Gran reported dense accumulations of diatoms adhering to the undersurface of during studies around 1900, attributing the brownish discoloration—known as "brown ice"—to high algal visible upon ice coring or breakage. Similar visual cues of ice staining were noted in Antarctic fast ice by explorers, with the 1901–1904 British National Antarctic Expedition (Discovery) collecting samples revealing cyanobacterial and communities contributing to ice pigmentation. These qualitative assessments, often opportunistic during navigation or overwintering, highlighted seasonal blooms triggered by spring penetration through thinning cover, with algal layers reaching thicknesses of several millimeters at ice-water interfaces. Early quantitative field efforts, emerging in the mid-20th century, built on these observations by measuring and . During the 1950s , Soviet and Norwegian teams in the documented chlorophyll concentrations exceeding 100 mg/m² in bottom ice layers via core sampling, linking algal growth to brine channel networks for nutrient and light access. In , U.S. expeditions in the 1950s–1960s confirmed comparable dynamics, with ice algae exhibiting photosynthetic rates up to 900 mg C/m²/day in under coastal fast ice, underscoring their role in extending polar primary production beyond open water seasons. Such data, derived from direct incubation and spectroscopic methods, refuted earlier dismissals of ice biota as mere transients, establishing foundational evidence for sympagic ecosystems despite challenges like ice instability and extreme salinity gradients.

Recent Empirical Findings (2023–2025)

In 2023, laboratory and field experiments conducted in revealed that sea-ice algal communities undergo pronounced seasonal physiological adjustments, remaining dormant during winter under low temperatures and high but rapidly activating metabolic processes in spring as conditions ameliorate. These studies measured photosynthetic rates and accumulation, demonstrating that in fast ice layers respond dynamically to gradients from brine rejection during ice formation, with peak productivity occurring in surface layers during melt onset. A satellite-based using CryoSat-2 altimetry data provided the first pan-Arctic estimates of potential onset timing, correlating earlier bloom initiation with regions of thinner ice and reduced snow cover, such as the central where blooms began up to 20-30 days prior to historical averages in low-ice years. This empirical mapping highlighted spatial variability, with bloom potentials advancing by 1-2 weeks per decade in marginal ice zones due to observed ice thinning trends from 2010-2023. In early 2025, observations in the Dalton Gulf, , documented high algal biomass incorporation into newly formed via dynamics, where turbulent freezing processes entrained from underlying waters, resulting in algal concentrations exceeding 10^6 cells per liter within days of formation and seeding subsequent under- blooms. Complementary experiments in fast exposed communities to varied light regimes, showing biomass migration toward upper layers (3-12 cm depth) under increased transmittance, with chlorophyll-a levels rising by factors of 2-5 in high-light treatments compared to controls. Mid-2025 Arctic expeditions identified thriving under-ice algal assemblages, including diatom-dominated communities beneath consolidated pack ice, with productivity rates estimated at 10-50 mg C m^{-2} day^{-1} in low-light conditions transmitted through thin , suggesting prior underestimations of basal production by up to 30% in models. These findings, derived from sampling and nutrient profiling, indicated by associated microbes as a key limiter, potentially amplifying carbon drawdown as cover diminishes.

Controversies and Unresolved Questions

Reliability of Biomarkers for Past Ice Conditions

Biomarkers such as the C25 highly branched isoprenoid (HBI) IP25, produced primarily by sympagic diatoms like Haslea kjellmanii within , serve as proxies for reconstructing past seasonal conditions through their preservation in marine . The presence of IP25 in sediment cores indicates historical cover, as its production is tied to ice algal growth during periods of sufficient penetration in spring, with concentrations typically highest near modern margins. Complementary indices like PIP25, which incorporate IP25 alongside open-water biomarkers such as brassicasterol, enable semi-quantitative assessments of extent, with values approaching 1 signaling and lower values reflecting variable or marginal conditions. Despite these strengths, the reliability of IP25-based reconstructions is constrained by spatial variability in ice algal production; for instance, IP25 is often absent in central sediments due to and limitations that suppress blooms, potentially leading to underestimation of past ice in such regions. Analytical challenges further complicate interpretations, including inconsistencies in measurement protocols across laboratories, such as differences in extraction methods and quantification standards, which can yield variable IP25 concentrations from the same samples by factors of up to 2–3 times. Zero IP25 values, common in open-water settings, introduce statistical biases in quantitative models like Bayesian calibrations, as they may reflect either ice absence or non-detection rather than true open conditions. In Antarctic contexts, analogous HBIs like IPSO25 from taxa such as Fragilariopsis curta show promise for reconstructing , correlating with modern pack ice edges, but face similar limitations including potential of biomarkers beyond ice margins and degradation during . Post-depositional processes, such as focusing and bioturbation, can homogenize signals over timescales of centuries, reducing resolution for high-frequency variability, while regional productivity gradients—driven by factors like stratification—may confound ice-specific interpretations without multi-proxy validation. Recent empirical calibrations, including 2024–2025 surface surveys, affirm moderate reliability for broad-scale reconstructions (e.g., glacial-interglacial shifts) but highlight the need for site-specific tuning to account for ecological variability, as evidenced by northward-decreasing sympagic signals beyond 73°N. Overall, while IP25 and related biomarkers provide robust qualitative evidence of past presence when detected above threshold levels (typically >1 ng/g), their quantitative reliability remains semi-empirical, with uncertainties estimated at 20–50% for extent proxies due to unmodeled factors like algal flux seasonality and pelagic contamination. Integration with independent proxies, such as dinocyst assemblages or isotopes, is essential to mitigate these gaps and enhance in paleoenvironmental studies.

Potential for Harmful Blooms and Geoengineering Relevance

Ice algae, primarily consisting of diatom species such as Nitzschia frigida and Fragilariopsis cylindrus, do not produce toxins themselves and are not classified as harmful algal bloom (HAB) species in the traditional sense of causing paralytic shellfish poisoning or direct neurotoxicity. However, their massive seasonal accumulations—reaching chlorophyll a concentrations up to 500 mg m⁻² in bottom ice communities—can contribute indirectly to ecological disruptions upon sea ice melt. These blooms release organic matter and nutrients into the underlying water column, potentially seeding post-melt phytoplankton proliferations that favor the expansion of toxic dinoflagellates like Alexandrium spp., which have been documented in recurrent Arctic blooms since 2012, with cell densities exceeding 10⁶ cells L⁻¹ in regions of recent ice loss. Observed increases in algal toxins, such as domoic acid and saxitoxins, in Arctic food webs correlate with reduced ice cover, as evidenced by elevated levels in bowhead whale feces from 2019–2023 sampling, linking ice melt-driven nutrient pulses to bioaccumulation risks for marine mammals and subsistence-harvested species. While direct causation remains correlative rather than experimentally proven, modeling and field data indicate that ice algae-derived carbon export—estimated at 15–50% of annual in ice-covered regions—exacerbates hypoxia in melt-influenced shelf waters, potentially amplifying secondary HAB risks from thriving in warmer, fresher surface layers. No widespread kills or incidents have been attributed solely to ice algae releases as of 2025, but monitoring programs in the Alaskan report a 20–30% rise in detections since 2018, underscoring an emerging threat as ice-free periods lengthen. In geoengineering contexts, ice algae's role in reducing —lowering reflectivity by 10–20% through pigmentation and accumulation—positions them as a target for interventions aimed at preserving polar ice sheets. Proposed strategies, such as deploying high- materials like hollow glass microspheres or artificial , could delay melt onset and thicken cover, thereby limiting light penetration to under-ice communities and suppressing algal blooms by 30–50% in simulations. Such modifications might mitigate feedback loops but risk altering biogeochemical cycles, as reduced ice algal production—historically contributing 10–25 Gt C year⁻¹ to —could diminish export fluxes to deeper waters, potentially shifting ecosystems toward pelagic dominance with unpredictable HAB dynamics. Empirical tests remain limited, with no large-scale deployments by , highlighting unresolved uncertainties in scaling these approaches without unintended trophic disruptions.

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