Protists in Arctic drift and land‐fast sea ice

Global climate change is having profound impacts on polar ice with changes in the duration and extent of both land‐fast ice and drift ice, which is part of the polar ice pack. Sea ice is a distinct habitat and the morphologically identifiable sympagic community living within sea ice can be readily distinguished from pelagic species. Sympagic metazoa and diatoms have been studied extensively since they can be identified using microscopy techniques. However, non‐diatom eukaryotic cells living in ice have received much less attention despite taxa such as the dinoflagellate Polarella and the cercozoan Cryothecomonas being isolated from sea ice. Other small flagellates have also been reported, suggesting complex microbial food webs. Since smaller flagellates are fragile, often poorly preserved, and are difficult for non‐experts to identify, we applied high throughput tag sequencing of the V4 region of the 18S rRNA gene to investigate the eukaryotic microbiome within the ice. The sea ice communities were diverse (190 taxa) and included many heterotrophic and mixotrophic species. Dinoflagellates (43 taxa), diatoms (29 taxa) and cercozoans (12 taxa) accounted for ~80% of the sequences. The sympagic communities living within drift ice and land‐fast ice harbored taxonomically distinct communities and we highlight specific taxa of dinoflagellates and diatoms that may be indicators of land‐fast and drift ice.     

Pan‐Arctic sea ice‐algal chl a biomass and suitable habitat are largely underestimated for multiyear ice

There is mounting evidence that multiyear ice (MYI) is a unique component of the Arctic Ocean and may play a more important ecological role than previously assumed. This study improves our understanding of the potential of MYI as a suitable habitat for sea ice algae on a pan‐Arctic scale. We sampled sea ice cores from MYI and first‐year sea ice (FYI) within the Lincoln Sea during four consecutive spring seasons. This included four MYI hummocks with a mean chl a biomass of 2.0 mg/m², a value significantly higher than FYI and MYI refrozen ponds. Our results support the hypothesis that MYI hummocks can host substantial ice‐algal biomass and represent a reliable ice‐algal habitat due to the (quasi‐) permanent low‐snow surface of these features. We identified an ice‐algal habitat threshold value for calculated light transmittance of 0.014%. Ice classes and coverage of suitable ice‐algal habitat were determined from snow and ice surveys. These ice classes and associated coverage of suitable habitat were applied to pan‐Arctic CryoSat‐2 snow and ice thickness data products. This habitat classification accounted for the variability of the snow and ice properties and showed an areal coverage of suitable ice‐algal habitat within the MYI‐covered region of 0.54 million km² (8.5% of total ice area). This is 27 times greater than the areal coverage of 0.02 million km² (0.3% of total ice area) determined using the conventional block‐model classification, which assigns single‐parameter values to each grid cell and does not account for subgrid cell variability. This emphasizes the importance of accounting for variable snow and ice conditions in all sea ice studies. Furthermore, our results indicate the loss of MYI will also mean the loss of reliable ice‐algal habitat during spring when food is sparse and many organisms depend on ice‐algae.     

Spring time production of bottom ice algae in the landfast sea ice zone at Barrow, Alaska

The primary production of bottom ice algae is an important food source for sympagic, pelagic and benthic organisms in the Arctic Ocean as well as Antarctic Ocean. Using ¹³C-¹⁵N isotope tracers, the recent ice algal production at Barrow during the spring season was lower in 2003 than three decades ago, although the maximum chlorophyll-a concentration for the bottom ice algae was similar to the values from previous studies. Estimated recent new and total production rates of the ice algae were 0.8 g C m^- 2 yr^- 1 and 2.0 g C m^- 2 yr^- 1 respectively, while the rates of water column phytoplankton were 0.2 g C m^- 2 yr^- 1 and 0.7 g C m^- 2 yr^- 1 for the spring sampling period in 2003. The ice algae contributed 74% of the pelagic primary production under the landfast sea ice at Barrow before the phytoplankton spring bloom. At the end of the season in 2003, a high carbon allocation of lipids in the ice algae was found. Three possible explanations- nutrient depletion, increasing light, and/or changes in species composition- were suggested for the high carbon incorporation into lipids. This high lipid synthesis of the bottom ice algae might be significant to zooplankton and benthic fauna grazers because lipids are the most energy dense biomolecules.     

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.     

Resource partitioning between Pacific walruses and bearded seals in the Alaska Arctic and sub-Arctic

Climate-mediated changes in the phenology of Arctic sea ice and primary production may alter benthic food webs that sustain populations of Pacific walruses (Odobenus rosmarus divergens) and bearded seals (Erignathus barbatus). Interspecific resource competition could place an additional strain on ice-associated marine mammals already facing loss of sea ice habitat. Using fatty acid (FA) profiles, FA trophic markers, and FA stable carbon isotope analyses, we found that walruses and bearded seals partitioned food resources in 2009–2011. Interspecific differences in FA profiles were largely driven by variation in non-methylene FAs, which are markers of benthic invertebrate prey taxa, indicating varying consumption of specific benthic prey. We used Bayesian multi-source FA stable isotope mixing models to estimate the proportional contributions of particulate organic matter (POM) from sympagic (ice algal), pelagic, and benthic sources to these apex predators. Proportional contributions of FAs to walruses and bearded seals from benthic POM sources were high [44 (17–67)% and 62 (38–83)%, respectively] relative to other sources of POM. Walruses also obtained considerable contributions of FAs from pelagic POM sources [51 (32–73)%]. Comparison of δ¹³C values of algal FAs from walruses and bearded seals to those from benthic prey from different feeding groups from the Chukchi and Bering seas revealed that different trophic pathways sustained walruses and bearded seals. Our findings suggest that (1) resource partitioning may mitigate interspecific competition, and (2) climate change impacts on Arctic food webs may elicit species-specific responses in these high trophic level consumers.     

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.     

Multiple ice‐binding proteins of probable prokaryotic origin in an Antarctic lake alga,Chlamydomonas sp. ICE‐MDV (Chlorophyceae)

Ice‐associated algae produce ice‐binding proteins (IBPs) to prevent freezing damage. The IBPs of the three chlorophytes that have been examined so far share little similarity across species, making it likely that they were acquired by horizontal gene transfer (HGT). To clarify the importance and source of IBPs in chlorophytes, we sequenced the IBP genes of another Antarctic chlorophyte, Chlamydomonas sp. ICE‐MDV (Chlamy‐ICE). Genomic DNA and total RNA were sequenced and screened for known ice‐associated genes. Chlamy‐ICE has as many as 50 IBP isoforms, indicating that they have an important role in survival. The IBPs are of the DUF3494 type and have similar exon structures. The DUF3494 sequences are much more closely related to prokaryotic sequences than they are to sequences in other chlorophytes, and the chlorophyte IBP and ribosomal 18S phylogenies are dissimilar. The multiple IBP isoforms found in Chlamy‐ICE and other algae may allow the algae to adapt to a greater variety of ice conditions than prokaryotes, which typically have a single IBP gene. The predicted structure of the DUF3494 domain has an ice‐binding face with an orderly array of hydrophilic side chains. The results indicate that Chlamy‐ICE acquired its IBP genes by HGT in a single event. The acquisitions of IBP genes by this and other species of Antarctic algae by HGT appear to be key evolutionary events that allowed algae to extend their ranges into polar environments.     

THE SHORT‐TERM EFFECT OF IRRADIANCE ON THE PHOTOSYNTHETIC PROPERTIES OF ANTARCTIC FAST‐ICE MICROALGAL COMMUNITIES1

Although sea‐ice represents a harsh physicochemical environment with steep gradients in temperature, light, and salinity, diverse microbial communities are present within the ice matrix. We describe here the photosynthetic responses of sea‐ice microalgae to varying irradiances. Rapid light curves (RLCs) were generated using pulse amplitude fluorometry and used to derive photosynthetic yield (Φ_PSII), photosynthetic efficiency (α), and the irradiance (E_k) at which relative electron transport rate (rETR) saturates. Surface brine algae from near the surface and bottom‐ice algae were exposed to a range of irradiances from 7 to 262 μmol photons · m^?2 · s^?1. In surface brine algae, Φ_PSII and α remained constant at all irradiances, and rETR_max peaked at 151 μmol photons · m^?2 · s^?1, indicating these algae are well acclimated to the irradiances to which they are normally exposed. In contrast, Φ_PSII, α, and rETR_max in bottom‐ice algae reduced when exposed to irradiances >26 μmol photons · m^?2 · s^?1, indicating a high degree of shade acclimation. In addition, the previous light history had no significant effect on the photosynthetic capacity of bottom‐ice algae whether cells were gradually exposed to target irradiances over a 12 h period or were exposed immediately (light shocked). These findings indicate that bottom‐ice algae are photoinhibited in a dose‐dependent manner, while surface brine algae tolerate higher irradiances. Our study shows that sea‐ice algae are able to adjust to changes in irradiance rapidly, and this ability to acclimate may facilitate survival and subsequent long‐term acclimation to the postmelt light regime of the Southern Ocean.     

Global change effects on the long‐term feeding ecology and contaminant exposures of East Greenland polar bears

Rapid climate changes are occurring in the Arctic, with substantial repercussions for arctic ecosystems. It is challenging to assess ecosystem changes in remote polar environments, but one successful approach has entailed monitoring the diets of upper trophic level consumers. Quantitative fatty acid signature analysis (QFASA) and fatty acid carbon isotope (δ¹³C‐FA) patterns were used to assess diets of East Greenland (EG) polar bears (Ursus maritimus) (n = 310) over the past three decades. QFASA‐generated diet estimates indicated that, on average, EG bears mainly consumed arctic ringed seals (47.5 ± 2.1%), migratory subarctic harp (30.6 ± 1.5%) and hooded (16.7 ± 1.3%) seals and rarely, if ever, consumed bearded seals, narwhals or walruses. Ringed seal consumption declined by 14%/decade over 28 years (90.1 ± 2.5% in 1984 to 33.9 ± 11.1% in 2011). Hooded seal consumption increased by 9.5%/decade (0.0 ± 0.0% in 1984 to 25.9 ± 9.1% in 2011). This increase may include harp seal, since hooded and harp seal FA signatures were not as well differentiated relative to other prey species. Declining δ¹³C‐FA ratios supported shifts from more nearshore/benthic/ice‐associated prey to more offshore/pelagic/open‐water‐associated prey, consistent with diet estimates. Increased hooded seal and decreased ringed seal consumption occurred during years when the North Atlantic Oscillation (NAO) was lower. Thus, periods with warmer temperatures and less sea ice were associated with more subarctic and less arctic seal species consumption. These changes in the relative abundance, accessibility, or distribution of arctic and subarctic marine mammals may have health consequences for EG polar bears. For example, the diet change resulted in consistently slower temporal declines in adipose levels of legacy persistent organic pollutants, as the subarctic seals have higher contaminant burdens than arctic seals. Overall, considerable changes are occurring in the EG marine ecosystem, with consequences for contaminant dynamics.     

Chlamydomonas carries out fatty acid β‐oxidation in ancestral peroxisomes using a bona fide acyl‐CoA oxidase

Peroxisomes are thought to have played a key role in the evolution of metabolic networks of photosynthetic organisms by connecting oxidative and biosynthetic routes operating in different compartments. While the various oxidative pathways operating in the peroxisomes of higher plants are fairly well characterized, the reactions present in the primitive peroxisomes (microbodies) of algae are poorly understood. Screening of a Chlamydomonas insertional mutant library identified a strain strongly impaired in oil remobilization and defective in Cre05.g232002 (CrACX2), a gene encoding a member of the acyl‐CoA oxidase/dehydrogenase superfamily. The purified recombinant CrACX2 expressed in Escherichia coli catalyzed the oxidation of fatty acyl‐CoAs into trans‐2‐enoyl‐CoA and produced H₂O₂. This result demonstrated that CrACX2 is a genuine acyl‐CoA oxidase, which is responsible for the first step of the peroxisomal fatty acid (FA) β‐oxidation spiral. A fluorescent protein‐tagging study pointed to a peroxisomal location of CrACX2. The importance of peroxisomal FA β‐oxidation in algal physiology was shown by the impact of the mutation on FA turnover during day/night cycles. Moreover, under nitrogen depletion the mutant accumulated 20% more oil than the wild type, illustrating the potential of β‐oxidation mutants for algal biotechnology. This study provides experimental evidence that a plant‐type FA β‐oxidation involving H₂O₂‐producing acyl‐CoA oxidation activity has already evolved in the microbodies of the unicellular green alga Chlamydomonas reinhardtii.     

Influence of sea ice on the composition of the spring phytoplankton bloom in the northern Baltic Sea

During the late winter and spring of 1994, the influence of sea ice on phytoplankton succession in the water was studied at a coastal station in the northern Baltic Sea. Ice cores were taken together with water samples from the underlying water and analysed for algal composition, chlorophyll a and nutrients. Sediment traps were placed under the ice and near the bottom, and the sedimented material was analysed for algal composition. The highest concentration of ice algae (4.1 mmol C m⁻²) was found shortly before ice break-up in the middle of April, coincidental with the onset of an under-ice phytoplankton bloom. The ice algae were dominated by the diatoms Chaetoceros wighamii Brightwell, Melosira arctica (Ehrenberg) Dickie and Nitzschia frigida Grunow. Under the ice the diatom Achnanthes taeniata Grunow and the dinoflagellate Peridiniella catenata (Levander) Balech were dominant. Calculations of sinking rates and residence times of the dominant ice algal species in the photic water column indicated that only one ice algal species (Chaetoceros wighamii) had a seeding effect on the water column: this diatom dominated the spring phytoplankton bloom in the water together with Achnanthes taeniata and Peridiniella catenata. Received: 9 May 1997 / Accepted: 15 February 1998     

Biogeochemistry and microbial community composition in sea ice and underlying seawater off East Antarctica during early spring

Pack ice, brines and seawaters were sampled in October 2003 in the East Antarctic sector to investigate the structure of the microbial communities (algae, bacteria and protozoa) in relation to the associated physico-chemical conditions (ice structure, temperature, salinity, inorganic nutrients, chlorophyll a and organic matter). Ice cover ranged between 0.3 and 0.8 m, composed of granular and columnar ice. The brine volume fractions sharply increased above −4°C in the bottom ice, coinciding with an important increase of algal biomass (up to 3.9 mg C l⁻¹), suggesting a control of the algae growth by the space availability at that period of time. Large accumulation of NH₄ ⁺ and PO₄ ³⁻ was observed in the bottom ice. The high pool of organic matter, especially of transparent exopolymeric particles, likely led to nutrients retention and limitation of the protozoa grazing pressure, inducing therefore an algal accumulation. In contrast, the heterotrophs dominated in the underlying seawaters.     

Fatty acid and stable isotope characteristics of sea ice and pelagic particulate organic matter in the Bering Sea: tools for estimating sea ice algal contribution to Arctic food web production

We determined fatty acid (FA) profiles and carbon stable isotopic composition of individual FAs (δ¹³C_FA values) from sea ice particulate organic matter (i-POM) and pelagic POM (p-POM) in the Bering Sea during maximum ice extent, ice melt, and ice-free conditions in 2010. Based on FA biomarkers, differences in relative composition of diatoms, dinoflagellates, and bacteria were inferred for i-POM versus p-POM and for seasonal succession stages in p-POM. Proportions of diatom markers were higher in i-POM (16:4n-1, 6.6–8.7 %; 20:5n-3, 19.6–25.9 %) than in p-POM (16:4n-1, 1.2–4.0 %; 20:5n-3, 5.5–14.0 %). The dinoflagellate marker 22:6n-3/20:5n-3 was highest in p-POM. Bacterial FA concentration was higher in the bottom 1 cm of sea ice (14–245 μg L^?1) than in the water column (0.6–1.7 μg L^?1). Many i-POM δ¹³C_FA values were higher (up to ~10 ‰) than those of p-POM, and i-POM δ¹³C_FA values increased with day length. The higher i-POM δ¹³C_FA values are most likely related to the reduced dissolved inorganic carbon (DIC) availability within the semi-closed sea ice brine channel system. Based on a modified Rayleigh equation, the fraction of sea ice DIC fixed in i-POM ranged from 12 to 73 %, implying that carbon was not limiting for primary productivity in the sympagic habitat. These differences in FA composition and δ¹³C_FA values between i-POM and p-POM will aid efforts to track the proportional contribution of sea ice algal carbon to higher trophic levels in the Bering Sea and likely other Arctic seas.     

Growth rates of arctic juvenile Scolelepis squamata (Polychaeta: Spionidae) isolated from Chukchi Sea fast ice

In spring, Arctic coastal fast ice is inhabited by high densities of sea ice algae and, among other fauna, juveniles of benthic polychaetes. This paper investigates the hypothesis that growth rates of juveniles of the common sympagic polychaete, Scolelepis squamata (Polychaeta: Spionidae), are significantly faster at sea ice algal bloom concentrations compared to concurrent phytoplankton concentrations. Juvenile S. squamata from fast ice off Barrow, Alaska, were fed with different algal concentrations at 0 and 5?°C, simulating ambient high sea ice algal concentrations, concurrent low phytoplankton concentrations, and an intermediate concentration. Growth rates, calculated using a simple linear regression equation, were significantly higher (up to 115?times) at the highest algal concentration compared to the lowest. At the highest algal concentration, juveniles grew faster at 5?°C compared to those feeding at 0?°C with a Q ₁₀ of 2.0. We conclude that highly concentrated sea ice algae can sustain faster growth rates of polychaete juveniles compared to the less dense spring phytoplankton concentrations. The earlier melt of Arctic sea ice predicted with climate change might cause a mismatch between occurrence of polychaete juveniles and food availability in the near future. Our data indicate that this reduction in food availability might counteract any faster growth of a pelagic juvenile stage based on forecasted increased water temperatures.     

Tracing carbon flow in an arctic marine food web using fatty acid-stable isotope analysis

Global warming and the loss of sea ice threaten to alter patterns of productivity in arctic marine ecosystems because of a likely decline in primary productivity by sea ice algae. Estimates of the contribution of ice algae to total primary production range widely, from just 3 to >50%, and the importance of ice algae to higher trophic levels remains unknown. To help answer this question, we investigated a novel approach to food web studies by combining the two established methods of stable isotope analysis and fatty acid (FA) analysis--we determined the C isotopic composition of individual diatom FA and traced these biomarkers in consumers. Samples were collected near Barrow, Alaska and included ice algae, pelagic phytoplankton, zooplankton, fish, seabirds, pinnipeds and cetaceans. Ice algae and pelagic phytoplankton had distinctive overall FA signatures and clear differences in delta(13)C for two specific diatom FA biomarkers: 16:4n-1 (-24.0+/-2.4 and -30.7+/-0.8 per thousand, respectively) and 20:5n-3 (-18.3+/-2.0 and -26.9+/-0.7 per thousand, respectively). Nearly all delta(13)C values of these two FA in consumers fell between the two stable isotopic end members. A mass balance equation indicated that FA material derived from ice algae, compared to pelagic diatoms, averaged 71% (44-107%) in consumers based on delta(13)C values of 16:4n-1, but only 24% (0-61%) based on 20:5n-3. Our estimates derived from 16:4n-1, which is produced only by diatoms, probably best represented the contribution of ice algae relative to pelagic diatoms. However, many types of algae produce 20:5n-3, so the lower value derived from it likely represented a more realistic estimate of the proportion of ice algae material relative to all other types of phytoplankton. These preliminary results demonstrate the potential value of compound-specific isotope analysis of marine lipids to trace C flow through marine food webs and provide a foundation for future work.     

Sourcing fatty acids to juvenile polar cod (Boreogadus saida) in the Beaufort Sea using compound-specific stable carbon isotope analyses

Juvenile polar cod (Boreogadus saida) are often found in close association with sea ice and represent an important trophic link in the Arctic food web. However, the proportional contribution of sea ice algal production via the sympagic food web to the diet of polar cod is unknown. To estimate the proportional contribution of fatty acids (FAs) from sea ice-derived particulate organic matter (i-POM) to the diet of juvenile polar cod, we used FA profiling and compound-specific stable carbon isotope analysis of individual FAs from juvenile polar cod collected from three regions in the Beaufort Sea. The δ¹³C values of the FAs 14:0, 16:4n-1, 18:0, 20:5n-3 and 22:6n-3 in the polar cod were found to most strongly resemble pelagic POM rather than i-POM. Results from isotope-mixing models using diatom FA markers indicated that the proportional contribution of FAs from i-POM to juvenile polar cod was ≤2 %, which suggests that juvenile polar cod had not sourced their FAs from i-POM. Thus, changes in sea ice coverage due to environmental change may not affect juvenile polar cod in regard to nutrients such as FAs but may still affect their populations by reducing critical shelter from predators.     

Diatom-Specific Highly Branched Isoprenoids as Biomarkers in Antarctic Consumers

The structure, functioning and dynamics of polar marine ecosystems are strongly influenced by the extent of sea ice. Ice algae and pelagic phytoplankton represent the primary sources of nutrition for higher trophic-level organisms in seasonally ice-covered areas, but their relative contributions to polar marine consumers remain largely unexplored. Here, we investigated the potential of diatom-specific lipid markers and highly branched isoprenoids (HBIs) for estimating the importance of these two carbon pools in an Antarctic pelagic ecosystem. Using GC-MS analysis, we studied HBI biomarkers in key marine species over three years in Adélie Land, Antarctica: euphausiids (ice krill Euphausia crystallorophias and Antarctic krill E. superba), fish (bald notothens Pagothenia borchgrevinki and Antarctic silverfish Pleuragramma antarcticum) and seabirds (Adélie penguins Pygoscelis adeliae, snow petrels Pagodroma nivea and cape petrels Daption capense). This study provides the first evidence of the incorporation of HBI lipids in Antarctic pelagic consumers. Specifically, a di-unsaturated HBI (diene) of sea ice origin was more abundant in ice-associated species than in pelagic species, whereas a tri-unsaturated HBI (triene) of phytoplanktonic origin was more abundant in pelagic species than in ice-associated species. Moreover, the relative abundances of diene and triene in seabird tissues and eggs were higher during a year of good sea ice conditions than in a year of poor ice conditions. In turn, the higher contribution of ice algal derived organic matter to the diet of seabirds was related to earlier breeding and higher breeding success. HBI biomarkers are a promising tool for estimating the contribution of organic matter derived from ice algae in pelagic consumers from Antarctica.     

Saving for the future: Pre‐winter uptake of algal lipids supports copepod egg production in spring

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Primary production of ice algae on a seasonally-ice-covered,continental shelf

Summary Ice algae samples were collected from the winter pack ice off Labrador during March 1984. The population was dominated by centric diatoms. Chlorophyll concentrations ranged from 40 to 190 mg m^–3, and particulate organic carbon from 2 to 10 g m^–3. Assimilation numbers for the ice algae ranged from 1.4 to 2.8 mg C (mg chl)^–1 h^–1 with a mean of 2.3, and were not significantly different from the assimilation numbers of the pelagic community beneath the ice. The ice algae were not photoinhibited at light intensities approaching surface light intensities. It is postulated that the dynamic nature of the ice field permits near-surface light intensities to reach the ice algae community at irregular intervals thereby suppressing photoinhibition.     

Freezing avoidance by supercooling in Olea europaea cultivars: the role of apoplastic water,solute content and cell wall rigidity

Plants can avoid freezing damage by preventing extracellular ice formation below the equilibrium freezing temperature (supercooling). We used Olea europaea cultivars to assess which traits contribute to avoid ice nucleation at sub‐zero temperatures. Seasonal leaf water relations, non‐structural carbohydrates, nitrogen and tissue damage and ice nucleation temperatures in different plant parts were determined in five cultivars growing in the Patagonian cold desert. Ice seeding in roots occurred at higher temperatures than in stems and leaves. Leaves of cold acclimated cultivars supercooled down to ?13 °C, substantially lower than the minimum air temperatures observed in the study site. During winter, leaf ice nucleation and leaf freezing damage (LT₅₀) occurred at similar temperatures, typical of plant tissues that supercool. Higher leaf density and cell wall rigidity were observed during winter, consistent with a substantial acclimation to sub‐zero temperatures. Larger supercooling capacity and lower LT₅₀ were observed in cold‐acclimated cultivars with higher osmotically active solute content, higher tissue elastic adjustments and lower apoplastic water. Irreversible leaf damage was only observed in laboratory experiments at very low temperatures, but not in the field. A comparative analysis of closely related plants avoids phylogenetic independence bias in a comparative study of adaptations to survive low temperatures.