Snow algae inhabit most of the cold regions worldwide, where long-lasting snow fields are common. The ecophysiology of snow algae has been studied intensively in North America and occasionally in polar regions. In the European Alps, the systematics of snow algae have been studied mainly by light microscopy. We studied temperature and light-dependence of photosynthesis, and plastid and extraplastid red pigment composition of red snow algae (Chlamydomonas nivalis) from snow patches in the high Alps of Austria. Both photosynthetic and respiratory data support the cryophilic adaptation of snow algal cells, but C. nivalis produced oxygen without any inhibition at temperatures up to 208C and maintained this for 1 h, at irradiances up to 1800 mmol m À2 s À1. Chlorophyll and primary carotenoid pigment composition was similar to that found in most other Chlorophyta. Additionally, large amounts of free and esterified astaxanthin were located in cytoplasmic lipid globules. Light and electron microscopy showed that the cell walls were frequently covered with tightly bound inorganic particles. Occasionally fungus-or bacteria-like structures were attached to the wall. The typical adult cell contained a single central chloroplast. Cytoplasmic structures were often difficult to resolve optically, as densely packed peripheral lipid globules, containing secondary carotenoids, occupied most of the cell volume. These pigments may shield the chloroplast from high irradiation (thus reducing the risk of photoinhibition) and may also be a potential carbon source during unfavourable climate conditions or the formation of daughter cells.
Surface ablation of the Greenland ice sheet is amplified by surface darkening caused by light‐absorbing impurities such as mineral dust, black carbon, and pigmented microbial cells. We present the first quantitative assessment of the microbial contribution to the ice sheet surface darkening, based on field measurements of surface reflectance and concentrations of light‐absorbing impurities, including pigmented algae, during the 2014 melt season in the southwestern part of the ice sheet. The impact of algae on bare ice darkening in the study area was greater than that of nonalgal impurities and yielded a net albedo reduction of 0.038 ± 0.0035 for each algal population doubling. We argue that algal growth is a crucial control of bare ice darkening, and incorporating the algal darkening effect will improve mass balance and sea level projections of the Greenland ice sheet and ice masses elsewhere.
Mesotaenium berggrenii is one of few autotrophs that thrive on bare glacier surfaces in alpine and polar regions. This extremophilic alga produces high amounts of a brownish vacuolar pigment, whose chemical constitution and ecological function is largely unknown until now. Field material was harvested to isolate and characterize this pigment. Its tannin nature was determined by photometric methods, and the structure determination was carried out by means of HPLC-MS and 1D- and 2D-NMR spectroscopy. The main constituent turned out to be purpurogallin carboxylic acid-6-O-β-d-glucopyranoside. This is the first report of such a phenolic compound in this group of algae. Because of its broad absorption capacities of harmful UV and excessive VIS radiation, this secondary metabolite seems to play an important role for the survival of this alga at exposed sites. Attributes and abundances of the purpurogallins found in M. berggrenii strongly suggest that they are of principal ecophysiological relevance like analogous protective pigments of other extremophilic microorganisms. To prove that M. berggrenii is a true psychrophile, photosynthesis measurements at ambient conditions were carried out. Sequencing of the 18S rRNA gene of this alpine species and of its arctic relative, the filamentous Ancylonema nordenskiöldii, underlined their distinct taxonomic position within the Zygnematophyceae.
Amongst a specialised group of psychrophilic microalgae that have adapted to thrive exclusively in summer snow fields, Chloromonas nivalis has been reported as a species causing green, orange or pink blooms in many alpine and polar regions worldwide. Nevertheless, the cytology, ecophysiology and taxonomy of this species are still unresolved. Intracellular processes during cyst formation, which is the dominant stage on snow fields, were examined with samples from the European Alps to better understand the cellular strategies of a green alga living in this harsh habitat. We show with two different methods, i.e. oxygen optode fluorometry and by chlorophyll fluorescence, that the cysts are photosynthetically highly active, although they do not divide, and that Chloromonas nivalis can cope with low as well as high light conditions. During cyst formation, the chloroplast is fragmented into several smaller parts, enlarging the surface to volume ratio. The pool of xanthophyll-cycle pigments is significantly enlarged, which is different from other snow algae. The cytoplasm is filled with lipid bodies containing astaxanthin, a secondary carotenoid that causes the typical orange colour. The cyst wall surface possesses characteristic elongate flanges, which are assembled extracellulary by accumulation of material in the periplasmatic interspace. Comparison of Chloromonas nivalis samples from different locations (Austrian Alps, Spitsbergen) by molecular methods indicates genetic variations due to spatial isolation, while a North American strain has no close relationship to the taxon.
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.
At the arctic archipelago of Svalbard, bare glacier surfaces are populated by microalgae like Ancylonema nordenskio¨ldii (Zygnematales, Streptophyta). The resulting blooms cause, due to a vacuolar pigmentation, brownish colourations of the glacier surface. This freshwater ice alga has been described from several polar and alpine glaciers; however, these reports lacked data about the ecophysiology or ultrastructure. Considering the harsh environmental conditions of the exceptional habitat, such as permanently low temperatures, exposure to high irradiation or a short vegetation period, the aim of this study was to elucidate cellular adaptations of A. nordenskio¨ldii. Thus, samples were collected at two glaciers in Spitsbergen. The cytoarchitecture of the cylindrical cells, which are arranged in unbranched filaments, demonstrates active cells with Golgi bodies, mitochondria and rough endoplasmic reticulum close to the nucleus when investigated by transmission electron microscopy (TEM). The cell walls are pore less and only 90 nm thin. A. nordenskio¨ldii only sporadically produces oblong zygotes when two filaments conjugate. The most remarkable cytological feature is peripheral brownish vacuoles, appearing osmiophil and electron dense by TEM. Aqueous extracts of this pigmentation show a broad absorption in the visible light and in the UV. Consequently, a protection against excessive irradiation is provided. Photosynthesis measurements performed at different temperatures and light levels indicate that the metabolism is adapted to temperatures close to the freezing point as well as to high light conditions. Therefore, A. nordenskio¨ldii can be regarded as metabolically and cytological well adapted to live on glaciers.
Ultraviolet (UV) radiation has become an important stress factor in polar regions due to anthropogenically induced ozone depletion. Although extensive research has been conducted on adaptations of polar organisms to this stress factor, few studies have focused on semi-terrestrial algae so far, in spite of their apparent vulnerability. This study investigates the effect of UV on two semi-terrestrial arctic strains (B, G) and one Antarctic strain (E) of the green alga Zygnema, isolated from Arctic and Antarctic habitats. Isolates of Zygnema were exposed to experimentally enhanced UV A and B (predominant UV A) to photosynthetic active radiation (PAR) ratio. The pigment content, photosynthetic performance and ultrastructure were studied by means of high-performance liquid chromatography (HPLC), chlorophyll a fluorescence and transmission electron microscopy (TEM). In addition, phylogenetic relationships of the investigated strains were characterised using rbcL sequences, which determined that the Antarctic isolate (E) and one of the Arctic isolates (B) were closely related, while G is a distinct lineage. The production of protective phenolic compounds was confirmed in all of the tested strains by HPLC analysis for both controls and UV-exposed samples. Moreover, in strain E, the content of phenolics increased significantly (p = 0.001) after UV treatment. Simultaneously, the maximum quantum yield of photosystem II photochemistry significantly decreased in UV-exposed strains E and G (p < 0.001), showing a clear stress response. The phenolics were most probably stored at the cell periphery in vacuoles and cytoplasmic bodies that appear as electron-dense particles when observed by TEM after high-pressure freeze fixation. While two strains reacted moderately on UV exposure in their ultrastructure, in strain G, damage was found in chloroplasts and mitochondria. Plastidal pigments and xanthophyll cycle pigments were investigated by HPLC analysis; UV A- and UV B-exposed samples had a higher deepoxidation state as controls, particularly evident in strain B. The results indicate that phenolics are involved in UV protection of Zygnema and also revealed different responses to UV stress across the three strains, suggesting that other protection mechanisms may be involved in these organisms.
Melting snow fields populated by aplanozygotes of the genus Chloromonas (Chlamydomonadales, Chlorophyta) are found in polar and alpine habitats. In the High Tatra Mountains (Slovakia), cells causing blooms of brownish–red snow designated as Scotiella tatrae kol turned out to be genetically (18S, ITS1 and ITS2 rDNA, rbcL) very closely related to Chloromonas nivalis (Chodat) Hoham et Mullet from the Austrian Alps. Therefore, Sc. tatrae is transferred into the latter taxon and reduced to a subspecies as Cr. nivalis subsp. tatrae. Both exhibit a similar photosynthetic performance, thrive in similar habitats at open sites above timberline, but differ in astaxanthin accumulation and number of aplanozygote cell wall flanges. In a field sample of Cr. nivalis subsp. tatrae, polyunsaturated fatty acids formed nearly 50 % of total lipids, dominating in phospholipids and glycolipids. Cr. nivalis subsp. tatrae represents likely a variation of a common cryoflora species with distinct morphology.
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