Hydrurus‐related golden algae (Chrysophyceae) cause yellow snow in polar summer snowfields

In polar regions, melting snow fields can be occupied by striking blooms of chrysophycean algae, which cause yellowish slush during summer. Samples were harvested at King George Island (South Shetland Islands, Maritime Antarctica) and at Spitsbergen (Svalbard archipelago, High Arctic). The populations live in an ecological niche, where water‐logged snow provides a cold and ephemeral ecosystem, possibly securing the survival of psychrophilic populations through the summer. A physiological adaptation to low temperatures was shown by photosynthesis measurements. The analysis of soluble carbohydrates showed the occurrence of glycerol and sugars, which may play a role in protection against intracellular freezing. Although both populations were made of unicells with Ochromonas‐alike morphology, investigation by molecular methods (18S rDNA sequencing) revealed unexpectedly a very close relationship to the mountain‐river dwelling Hydrurus foetidus (Villars) Trevisan. However, macroscopic thalli typical for the latter species were never found in snow, but are known from nearby localities, and harvested samples of snow algae exposed to dryness evolved a similar pervading, ‘fishy’ smell. Moreover, in both habitats tetrahedal zoospores with four elongate spikes were found, similar to what is known from Hydrurus. Our molecular results go along with earlier reports, where chrysophycean sequences of the same taxonomic affiliation were isolated from snow. This points to a distinct group of photoautotrophic, Hydrurus‐related chrysophytes, which are characteristic for long‐lasting, slowly melting snow packs in certain cold regions of the world.     

Aggregation of algae released from melting sea ice: implications for seeding and sedimentation

Summary Factors influencing the fate of ice algae released from melting sea ice were studied during a R V Polarstern cruise (EPOS Leg 2) to the northwestern Weddell Sea. The large-scale phytoplankton distribution patterns across the receding ice edge and small-scale profiling of the water column adjacent to melting ice floes indicated marked patchiness on both scales. The contribution of typical ice algae to the phytoplankton was not significant. In experiments simulating the conditions during sea ice melting, ice algae revealed a strong propensity to form aggregates. Differences in the aggregation potential were found for algal assemblages collected from the ice interior and the infiltration layer. Although all algal species collected from the ice were also found in aggregates, the species composition of dispersed and aggregated algae differed significantly. Aggregates were of a characteristic structure consisting of monospecific microaggregates which are likely to have formed in the minute brine pockets and channels within the ice. Sinking rates of aggregates were three orders of magnitude higher than those of dispersed ice algae. These observations, combined with the negligible seeding effect of ice algae found during this study, suggest that ice algae released from the melting sea ice are subject to rapid sedimentation. High grazing pressure at the ice edge of the investigation area is another factor eliminating ice algae released during melting.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

PHOTOSYNTHESIS IN THE SNOW: THE ALGA CHLAMYDOMONAS NIVALIS (CHLOROPHYCEAE)1

Red blooms of snow algae consisting almost exclusively of large spherical red cells of Chlamydomonas nivalis (Bauer) Wille are widespread during the summer in the Beartooth Mountains in Montana and Wyoming. Field studies designed to examine the effects of temperature, light, and water potential on algal activity were performed with natural populations using photosynthetic ¹⁴C-HCO₃- or ¹⁴CO₂ incorporation as a measure of activity. The algae photo-synthesized optimally at 5.4 × 10⁴ lx, but were not inhibited by increased light intensity up to 8.6 × 10⁴ lx, the maximum observed in the field. Photosynthesis was sensitive to a reduction in water potential, and since low water potentials develop in snow at temperatures below 0 C, it is unlikely that significant algal activity occurs at the sub-0 temperatures which occur throughout winter. Photosynthesis was much lower following melting of the snow, but this was probably due to decreased diffusion of CO₂. The optimal temperatures varied considerably among the different algal populations. Most samples photo-synthesized optimally at 10 or 20 C but retained substantial activity at temperatures as low as 0 or -3 C. Exceptional samples photosynthesized optimally at 0 or -3 C. It is proposed that the varied temperature responses reflect the presence of different temperature strains. Taken together, the data suggest that development of the snow algae can occur only during the summer months.     

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.     

Light climate and phytoplankton photosynthesis in maritime Antarctic lakes

The responses of phytoplankton populations to seasonal changes in radiation flux in two Antarctic lakes with extensive winter ice-cover are described. A phytoplankton capable of photosynthesis was found throughout the year in both systems. During winter, low incident radiation combined with thick layers of snow and ice prevented in situ photosynthesis becoming detectable. The beginning of spring was marked by a reduction in snow cover which resulted in a considerable increase in surface penetrating radiation. Planktonic algae rapidly adapted to utilise these increased levels efficiently, though they still showed characteristics of strong shade adaptation.Loss of ice cover at the start of the short open water period further increased the radiation levels and a summer population developed which was much less shade adapted. Saturation and photoinhibition effects were widespread during this period as the algae proved unable to utilise high radiation levels efficiently. They were however effective at the radiation fluxes prevalent in the lower part of the rapidly circulating water columns.     

Snow and Glacial Algae: A Review1

Snow or glacial algae are found on all continents, and most species are in the Chlamydomonadales (Chlorophyta) and Zygnematales (Streptophyta). Other algal groups include euglenoids, cryptomonads, chrysophytes, dinoflagellates, and cyanobacteria. They may live under extreme conditions of temperatures near 0°C, high irradiance levels in open exposures, low irradiance levels under tree canopies or deep in snow, acidic pH, low conductivity, and desiccation after snow melt. These primary producers may color snow green, golden-brown, red, pink, orange, or purple-grey, and they are part of communities that include other eukaryotes, bacteria, archaea, viruses, and fungi. They are an important component of the global biosphere and carbon and water cycles. Life cycles in the Chlamydomonas–Chloromonas–Chlainomonas complex include migration of flagellates in liquid water and formation of resistant cysts, many of which were identified previously as other algae. Species differentiation has been updated through the use of metagenomics, lipidomics, high-throughput sequencing (HTS), multi-gene analysis, and ITS. Secondary metabolites (astaxanthin in snow algae and purpurogallin in glacial algae) protect chloroplasts and nuclei from damaging PAR and UV, and ice binding proteins (IBPs) and polyunsaturated fatty acids (PUFAs) reduce cell damage in subfreezing temperatures. Molecular phylogenies reveal that snow algae in the Chlamydomonas–Chloromonas complex have invaded the snow habitat at least twice, and some species are polyphyletic. Snow and glacial algae reduce albedo, accelerate the melt of snowpacks and glaciers, and are used to monitor climate change. Selected strains of these algae have potential for producing food or fuel products.     

OBSERVATIONS ON SNOW ALGAE IN CALIFORNIA1,2

Algae that impart a red color to snowfields are rather common in California. Red snow occurs mainly in the Sierra Nevada at altitudes of 10,000–12,000 ft (3050–3600 in) and can occur at high altitudes where snow persists in other parts of the state. The distribution in the Sierra was similar in 1969 and 1970, contrasting snowfall years. Colored snow was found from May to October in old, wet snow-fields. The predominant color was red and occurred as surface patches in depressions in the snow. The color could extend as deep as 30 cm below the snow surface. Algae in the snowfields of the Tioga Pass area (Sierra Nevada) were large, red, spherical cells of Chlamydomonas nivalis. No other algae were seen. Their distribution, as measured by cell numbers and chlorophyll a, was patchy. Algal cells and chlorophyll a were mainly distributed at or near the snow surface but extended down to a depth of 10 cm. Light intensity was greatly attenuated by snow, but enough light for photosynthesis was found at 50 cm below the surface. Nutrient content of one snow sample was very low. The populations were very actively photosynthetic and took up as much as 65% of added ¹⁴CO₂ in only 3 hr. It was tentatively concluded that CO₂ limits in situ photosynthesis. Photosynthesis was inhibited by melting snow samples. Rough calculations of the growth rate suggested in situ generation times of only a few days for these algae.     

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.     

Is Snow a sufficient Source of Water for Horses kept Outdoors in Winter? A Case Report

Due to extreme weather conditions, a flock of outwintered Icelandic horses had to manage for several days on snow as the source of free water. They were fed grass silage ad lib, and any change in feed consumption was not observed. After nine days, blood samples were taken and analysed for plasma osmolality, they were subjected to a simple clinical examination, and offered drinking water. Osmolality levels were within normal limits and mean value did not differ significantly from samples which previously were taken of the same individuals. The general condition of the horses was normal, with no signs of clinical dehydration or disease. The horses showed very little interest for the offered drinking water. This suggests that in cold winter weather, horses being fed grass silage and adjusted to eat snow, can manage for several days with snow substituting liquid water without their physiology and welfare being challenged.     

古尔班通古特沙漠南缘融雪水土壤入渗量

融雪水土壤入渗量是干旱区沙漠重要的水平衡收入项.2012-2013、2013-2014年两个冬季对古尔班通古特沙漠南缘沙丘西坡、东坡和丘间地降雪前和融雪后的土壤含水率进行监测,根据水量平衡原理计算了沙丘西坡、东坡、丘间地和景观尺度上的融雪水土壤入渗量,并与采用筒测法的实测结果进行比较.结果表明： 降雪前土壤含水率较低,未冻层非饱和土壤水对地表冻结层土壤水分的补给可忽略不计;融雪入渗水是表层土壤获得补给的主要水源;冻结期潜水既没有蒸发,积雪融化后潜水也没有获得补给;研究区西坡、东坡、丘间地和景观尺度上的融雪水土壤入渗量分别为20～43、27～43、32～45和26～45 mm.
     

Phylogeny and biogeography of an uncultured clade of snow chytrids

Numerous studies have shown that snow can contain a diverse array of algae known as ‘snow algae’. Some reports also indicate that parasites of algae (e.g. chytrids) are also found in snow, but efforts to phylogenetically identify ‘snow chytrids’ have not been successful. We used culture‐independent molecular approaches to phylogenetically identify chytrids that are common in long‐lived snowpacks of Colorado and Europe. The most remarkable finding of the present study was the discovery of a new clade of chytrids that has representatives in snowpacks of Colorado and Switzerland and cold sites in Nepal and France, but no representatives from warmer ecosystems. This new clade (‘Snow Clade 1’ or SC1) is as deeply divergent as its sister clade, the Lobulomycetales, and phylotypes of SC1 show significant (P < 0.003) genetic‐isolation by geographic distance patterns, perhaps indicating a long evolutionary history in the cryosphere. In addition to SC1, other snow chytrids were phylogenetically shown to be in the order Rhizophydiales, a group with known algal parasites and saprotrophs. We suggest that these newly discovered snow chytrids are important components of snow ecosystems where they contribute to snow food‐web dynamics and the release of nutrients due to their parasitic and saprotrophic activities.     

Community structure of microorganisms associated with reddish-brown iron-rich snow

Reddish-brown colored snow, containing spherical brown particles, has been observed in several mires in Japan. In order to characterize this remarkable phenomenon, the microbial community and chemical species in snow were analyzed. A core sample of snow which had a colored region was investigated and it revealed vertical shifts in physicochemical characteristics and the microbial community structure. The abundance of particles peaked within the colored layer, and correlated with the amount of reducible Fe(III). The interstitial water of the colored layer was enriched with Fe(II), and characterized by reduced concentration of dissolved methane. The bacterial community in the colored region was characterized by higher relative abundance of iron-reducing bacteria and methanotrophs. Aggregates of the brown particles were found as precipitates in snow melt pools, and were subjected to cloning analyses targeting several different genes. The majority of bacterial 16S rRNA gene clones belonged to the class Betaproteobacteria or the phylum Bacteroidetes. No snow algae were detected in the eukaryotic small subunit rRNA gene clone library. As a possible carbon source to sustain the community in the snow, involvements of carbon dioxide and methane were investigated by analyzing the genes involved in their assimilation. In the analyses of genes for ribulose-1,5-biphosphate carboxylase/oxygenase, clones related to sulfur oxidizers were obtained. The analysis of particulate methane monooxygenase genes indicated dominance of Methylobacter species. These results emphasized the uniqueness of this phenomenon, and iron reducers of the genus Geobacter are suggested to be the key organisms that could be investigated in order to understand the mechanism of this phenomenon.     

Snow algae from northwest Svalbard: their identification, distribution, pigment and nutrient content

We mapped coloured snow during the summers of 1995 and 1996 at about 60 localities in the coastal region of northwest Spitsbergen. The colour was mainly induced by snow algae (Chlamydomonas spp. and Chloromonas spp.). In the late summer of 1996, snow algal fields of several hundred meters in size were observed along the west and north coasts. They had no preferred geographical orientation. We studied the abundance of primary pigments and secondary carotenoids from different developmental stages of the snow algae of Chlamydomonas spp. under natural conditions. Extensive accumulation of astaxanthin and its esters accompanied the transition from green biflagellated cells to orange spores, hypnozygotes and dark-red cysts. The photoprotective effect of the secondary carotenoids is enhanced by concentration in cytoplasmic lipid droplets around the nucleus and chloroplast. The nutrient content of melt-water and snow algae had no direct correlation with the content of secondary carotenoids. Relatively high Fe, Ca, P, K and Al contents of snow algae were found, suggesting a good supply of these mineral elements. Received: 20 May 1997 / Accepted: 18 March 1998     

雪被去除对川西高山冷杉林冬季土壤微生物生物量碳氮和可培养微生物数量的影响

为了解气候变暖情景下雪被减少对冬季土壤微生物特征的影响,采用人工遮雪的方法,研究了雪被去除对原始冷杉林土壤微生物生物量和可培养微生物数量的影响.结果表明:雪被去除显著影响土壤微生物生物量碳(MBC)和氮(MBN)以及可培养细菌和真菌数量,但土壤微生物在雪被覆盖不同阶段具有不同的响应特征.在雪被去除处理下,土壤有机层MBC和MBN在雪被形成初期和雪被融化前期显著降低,而在雪被覆盖期和雪被融化后期显著增加;在雪被形成初期至雪被覆盖期,可培养细菌数量都显著降低,但可培养真菌数量都显著增加.雪被融化后,雪被去除显著降低土壤有机层MBC和可培养真菌数量,显著增加可培养细菌数量,对MBN无显著影响.矿质土壤层MBC、MBN和可培养微生物数量在雪被去除下的变化趋势与土壤有机层基本一致,但波动较小.雪被去除还改变了川西高山冷杉林冬季土壤微生物类群比,提高了土壤可培养真菌数量的冬季优势.     

Water loss and temperature interact to compound amphibian vulnerability to climate change

Ectotherm thermal physiology is frequently used to predict species responses to changing climates, but for amphibians, water loss may be of equal or greater importance. Using physical models, we estimated the frequency of exceeding the thermal optimum (T_opt) or critical evaporative water loss (EWL_crit) limits, with and without shade‐ or water‐seeking behaviours. Under current climatic conditions (2002–2012), we predict that harmful thermal (>T_opt) and hydric (>EWL_crit) conditions limit the activity of amphibians during ~70% of snow‐free days in sunny habitats. By the 2080s, we estimate that sunny and dry habitats will exceed one or both of these physiological limits during 95% of snow‐free days. Counterintuitively, we find that while wet environments eliminate the risk of critical EWL, they do not reduce the risk of exceeding T_opt (+2% higher). Similarly, while shaded dry environments lower the risk of exceeding T_opt, critical EWL limits are still exceeded during 63% of snow‐free days. Thus, no single environment that we evaluated can simultaneously reduce both physiological risks. When we forecast both temperature and EWL into the 2080s, both physiological thresholds are exceeded in all habitats during 48% of snow‐free days, suggesting that there may be limited opportunity for behaviour to ameliorate climate change. We conclude that temperature and water loss act synergistically, compounding the ecophysiological risk posed by climate change, as the combined effects are more severe than those predicted individually. Our results suggest that predictions of physiological risk posed by climate change that do not account for water loss in amphibians may be severely underestimated and that there may be limited scope for facultative behaviours to mediate rapidly changing environments.     

Microscopical aspects on symbiosis of Spongilla lacustris (Porifera,Spongillidae) and green algae

Summary When growing in the sunlight, some specimens of Spongilla lacustris are coloured green due to the presence of symbiotic unicellular chlorellae. The algae live inside most sponge cells. The chlorellae were extracted from green sponges, cultivated, added to algae-free sponges and fixed after different incubation times. In this way the uptake of the algae, their distribution and their final whereabouts in the mesenchymatic cells could be followed by in vivo microscopy, phase-contrast microscopy and electron microscopy. A few minutes after addition, the chlorellae can be found inside the choanocyte chambers. Here they are taken up by the cell bodies and collars of the choanocytes. Pinacocytes are also involved in the uptake. The distribution of algae results from a specific transmission from the donor cell to the receiver cell. The chlorellae are not released from their host vacuoles until they are extensively enclosed by the cell taking them up. Six hours after addition, all sponge cells contain algae except granulocytes, microscleroblasts, the pinacocytes of the peripheral rim region and those of the pinacoderm. The chlorellae are able to divide inside the sponge cells.Abbreviations StM Stereo-microscopical photograph - PhC Phase-contrast microscopical photograph - EM Electron microscopical photograph     

A model for the hydrological cycle of a forested catchment and assessment of the changes caused in water balance by cuttings

A physicomathematical model of the hydrological cycle in a forested catchment was constructed. This model describes the interception of liquid and solid precipitation by tree crowns; snow accumulation and melting; vertical transfer of moisture in soil and its evaporation; and surface, subsurface, and channeled runoffs. The model was calibrated and verified using the observation data for the completely forested Taezhnyi catchment within the area of the Valdai Water Balance Station. Then the model was used to assess possible changes in the hydrological cycle after clear cuttings in this catchment. The values of model parameters were compared to the corresponding soil characteristics in the adjacent treeless (field) Usad’evskii catchment. Modeling results demonstrate that the average water reserve in the snow cover before melting can increase by 15% after forest cutting in the Taezhnyi catchment. The losses for snow sublimation are reduced almost two-fold. The snow melting intensity increases by 30% and its duration decreases by 10 days. The annual runoff after cutting increases by 7–10%; however, the seasonal distribution of the runoff and the constituents of the water balance change to a greater degree. During spring flood, the maximal water discharge in the forested catchment is 50% smaller than after forest cutting. The duration of spring flood after cutting is reduced by 5–7 days. The changes in the hydrological cycle depending on the age-related alteration in leaf area index were also studied.     

The effect of water turbidity on the near-surface water temperature of larval habitats of the malaria mosquito <Emphasis Type="Italic">Anopheles gambiae</Emphasis>

Water temperature is an important determinant in many aquatic biological processes, including the growth and development of malaria mosquito (Anopheles arabiensis and A. gambiae) immatures. Water turbidity affects water temperature, as suspended particles in a water column absorb and scatter sunlight and hence determine the extinction of solar radiation. To get a better understanding of the relationship between water turbidity and water temperature, a series of semi-natural larval habitats (diameter 0.32 m, water depth 0.16 m) with increasing water turbidity was created. Here we show that at midday (1300 hours) the upper water layer (thickness of 10 mm) of the water pool with the highest turbidity was on average 2.8°C warmer than the same layer of the clearest water pool. Suspended soil particles increase the water temperature and furthermore change the temperature dynamics of small water collections during daytime, exposing malaria mosquito larvae, which live in the top water layer, longer to higher temperatures.     

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.