Large brown seaweeds (kelps) represent ecologically important habitats in coastal systems. Along the western shores of the Svalbard archipelago, the kelp Alaria esculenta forms dense stands and provides habitat to a multitude of associated organisms. At the growth site in shallow waters, kelp sporophytes can be exposed to high irradiances of harmful ultraviolet (UV-B) radiation resulting from a thinning of the stratospheric ozone layer over the Arctic. Here, we tested to what extent UV-B-induced impairment of photophysiology is accompanied by changes in the cellular ultrastructure of A. esculenta and characterizes sites of cellular damage after UV exposure. Adverse effects on cellular components mostly manifest in chloroplast thylakoid membrane integrity, which is accompanied by a decrease in maximal quantum yield of photosystem II. At the applied irradiances of UV radiation, A. esculenta is stressed to an intermediate extent confirming its adaptability to a moderately increased UV-B climate.

Due to modern life with increasing traffic, industrial production and agricultural practices, high amounts of heavy metals enter ecosystems and pollute soil and water. As a result, metals can be accumulated in plants and particularly in algae inhabiting peat bogs of low pH and high air humidity. In the present study, we investigated the impact and intracellular targets of aluminum, copper, cadmium, chromium VI and zinc on the filamentous green alga Desmidium swartzii, which is an important biomass producer in acid peat bogs. By means of transmission electron microscopy (TEM) and electron energy loss spectroscopy (EELS) it is shown that all metals examined are taken up into Desmidium readily, where they are sequestered in cell walls and/or intracellular compartments. They cause effects on cell ultrastructure to different degrees and additionally disturb photosynthetic activity and biomass production. Our study shows a clear correlation between toxicity of a metal and the ability of the algae to compartmentalize it intracellularly. Cadmium and chromium, which are not compartmentalized, exert the most toxic effects. In addition, this study shows that the filamentous alga Desmidium reacts more sensitively to aluminum and zinc when compared to its unicellular relative Micrasterias, indicating a severe threat to the ecosystem.

The effect of a wide range of temperatures (−15 and 60°C) in darkness or under strong irradiation [1,600 μmol(photon) m −2 s −1 ] on quantum yield of photosystem II photochemistry and xanthophyll cycle pigments was investigated in a tropical fruit crop ( Musa sp.) and a temperate spring flowering plant ( Allium ursinum L.). In darkness within the nonlethal thermal window of A. ursinum (from −6.7 to 47.7°C; 54.5 K) and of Musa sp. (from −2.2°C to 49.5°C; 51.7 K) maximal quantum yield of PSII photochemistry (F v /F m ) was fairly unaffected by temperature over more than 40 K. At low temperature F v /F m started to drop with ice nucleation but significantly only with initial frost injuries (temperature at 10% frost damage; LT 10 ). The critical high temperature threshold for PSII (T c ) was 43.8°C in A. ursinum and 44.7°C in Musa sp. Under strong irradiation, exposure to temperatures exceeding the growth ones but being still nonlethal caused photoinhibition in both species. Severity of photoinhibition increased with increasing distance to the growth temperature range. ΔF/F m ′ revealed distinctly different optimum temperature ranges: 27–36°C for Musa sp. and 18–27°C for A. ursinum exceeding maximum growth temperature by 2–7 K. In both species only at temperatures > 30°C zeaxanthin increased and violaxanthin decreased significantly. At nonlethal low temperature relative amounts of xanthophylls remained unchanged. At temperatures > 40°C β-carotene increased significantly in both species. In Musa sp. lutein and neoxanthin were significantly increased at 45°C, in A. ursinum lutein remained unchanged, neoxanthin levels decreased in the supraoptimal temperature range. In darkness, F v /F m was highly temperature-insensitive in both species. Under strong irradiation, whenever growth temperature was exceeded, photoinhibition occurred with xanthophylls being changed only under supraoptimal temperature conditions as an antiradical defence mechanism.

In Antarctica, mass accumulations of psychrophilic algae cause striking phenomena like green, orange, or red snow. This occurs during summer, when coastal snowfields start to melt, become waterlogged and photoautotrophs can thrive. Chloromonas polyptera (Fritsch) Hoham, Mullet & Roemer is a unicellular species that causes orange snow in the vicinity of penguin rockeries. It has been recognized for many decades because of the distinct habitat and the characteristic morphology of cysts with elongated flanges on the outer cell wall. However, closer investigations concerning the ecology or physiology have been sparse so far. Field material was collected from two sites on the Antarctic Peninsula to find out more about metabolic and cellular strategies. The results were compared with a closely related species from high alpine locations, Chloromonas nivalis (Chodat) Hoham & Mullet. Despite the geographical distance, C. polyptera shares several physiological strategies with the alpine relative, such as the formation of cyst stages, saccharose and glycerol as main soluble carbohydrates and the abundant accumulation of the carotenoid astaxanthin. Moreover, photosynthesis is adapted to temperatures of about 1 degrees C. The molecular phylogeny confirmed a close relationship of C. polyptera to other Chloromonas species isolated from snow. Chloromonas polyptera seems to be exclusive to coastal Antarctic ecosystems influenced by animal nutrient input.

At the arctic archipelago of Svalbard, bare glacier surfaces are populated by microalgae like Ancylonema nordenskiöldii (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. nordenskiöldii. 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. nordenskiöldii 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. nordenskiöldii can be regarded as metabolically and cytological well adapted to live on glaciers.

Klebsormidium crenulatum (Kütz.) Lokhorst (Klebsormidiophyceae, Streptophyta) isolated from an alpine soil in Tyrol, Austria, was experimentally exposed to desiccation under various relative air humidities (RH 5, 75, and >95%, ambient air 55%-60%). The effects on the structure and ultrastructure of K. crenulatum after 1, 4, or 7 d of desiccation at 5, 75, and >95% RH were investigated. The cross walls were deformed to an undulated shape, and the cell diameter was reduced to ∼60% of the control. Regardless of the RH applied, in all cases the cytoplasm appeared denser compared to that of liquid-culture-grown cells. Electron-dense particles with diameters of 0.4 μm-0.8 μm were observed in the cytoplasm, likely representing lipid droplets. The chloroplasts of desiccated samples contained a large number of plastoglobules. The number and appearance of mitochondria were not visibly altered, as also verified by 3,3' dihexyloxacarbocyanine iodine (DIOC6 ) staining. The amphiphilic styryl dye FM 1-43 resulted in staining of the plasma membrane in cells from liquid culture. In 7 d desiccated samples, a marked fluorescence is seen in ∼40%-50% of the cells, which were dead. Actin microfilaments (MFs) were drastically disrupted after desiccation; only dotlike actin batches remained. These results demonstrate that flexibility of the cell walls and maintenance of the key organelles play a key role in the tolerance of desiccation stress in K. crenulatum.

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.

An indoor sun simulator was used to provide elevated UV-B radiation (280-315 nm) in combination with realistic ratios to PAR (400-700 nm) and UV-A radiation (315-400 nm) in order to test the physiological response of a soil- and snow microalga during a three-day stress scenario, which may occasionally occur in their respective arctic and alpine habitats.Chlamydomonas nivalis and Tetracystis sp. are initial colonizers of harsh habitats like summer snow fields and bare arctic soils. The two species were chosen because of their role as primary successors in places where life is generally limited by extreme climatic and nutritional conditions.The influence of the increased UV-B irradiation (1.43 W m(-2); control: 0.52 W m(-2)) on photosynthesis and pigment composition was measured. Both species survived this incubation without any morphological signs of damage, but oxygen production was reduced by 20-56%. Under control conditions, the amount of chlorophylls (Chls) and carotenoids (Cars) per dry mass increased after three days due to optimal light conditions. After the same period, the treated samples of the soil alga Tetracystis sp. showed a smaller increase in Chls and primary Cars than the control. However, the production of extraplastidal, secondary Cars was induced. On the contrary, the snow alga C. nivalis already had high amounts of secondary Cars before the experiment, and after exposure, all pigment classes increased more compared to control conditions. The results show that these microalgae can tolerate short episodes of enhanced UV-B radiation. Photosynthesis may be temporally impaired, but the cells respond by the production of secondary Cars, which can shield their chloroplasts against excessive irradiation or quench reactive oxygen species.

The life of plants growing in cold extreme environments has been well investigated in terms of morphological, anatomical, and ecophysiological adaptations. In contrast, long-term cellular or metabolic studies have been performed by only a few groups. Moreover, a number of single reports exist, which often represent just a glimpse of plant behavior. The review draws together the literature which has focused on tissue and cellular adaptations mainly to low temperatures and high light. Most studies have been done with European alpine plants; comparably well studied are only two phanerogams found in the coastal Antarctic. Plant adaptation in northern polar regions has always been of interest in terms of ecophysiology and plant propagation, but nowadays, this interest extends to the effects of global warming. More recently, metabolic and cellular investigations have included cold and UV resistance mechanisms. Low-temperature stress resistance in plants from cold environments reflects the climate conditions at the growth sites. It is now a matter of molecular analyses to find the induced genes and their products such as chaperones or dehydrins responsible for this resistance. Development of plants under snow or pollen tube growth at 0°C shows that cell biology is needed to explain the stability and function of the cytoskeleton. Many results in this field are based on laboratory studies, but several publications show that it is not difficult to study cellular mechanisms with the plants adapted to a natural stress. Studies on high light and UV loads may be split in two parts. Many reports describe natural UV as harmful for the plants, but these studies were mainly conducted by shielding off natural UV (as controls). Other experiments apply additional UV in the field and have had practically no negative impact on metabolism. The latter group is supported by the observations that green overwintering plants increase their flavonoids under snow even in the absence of UV. Thus, their defense and antioxidant role dominates. Ultrastructural comparisons were unable to find special light adaptations in plants taken from polar regions vs. high alpine species. The only adaptation found at the subcellular level for most alpine and polar plants are protrusions of the chloroplast envelopes. They are seen as a demand for fast membrane transport requiring additional membrane surface area, whereby the increase in stroma volume may help to support carbohydrate formation. Plants forming such protrusions have to cope with a short vegetation time. These observations are connected to the question as to how photosynthesis works quite well even at or under zero temperatures. The interplay between plastids, mitochondria, and peroxisomes, known as photorespiration, seems to be more intense than in lowland plants. This organelle cooperation serves as a valve for a surplus in solar energy input under cold conditions. Additional metabolic acclimations are under investigation, such as the role of an alternative plastid terminal oxidase. Plants from cold environments may also be seen as ideal objects for studying the combined effects of high light plus cold resistance—from the molecular level to the whole plant adaptation. Modern instrumentation makes it possible to perform vital metabolic measurements under outdoor conditions, and research stations in remote polar and alpine areas provide support for scientists in the preparation of samples for later cellular studies in the home laboratory.

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.

Der typische Abbau von Chlorophyll in alternden Blättern führt zu farblosen und schwer detektierbaren Kataboliten. Wie B. Kräutler et al. in ihrer Zuschrift auf S. 5300 ff. berichten, häufen sich aber blau fluoreszierende Chlorophyll-Kataboliten in gelb werdenden Bananenblättern an, die dann unter UV-Licht blau lumineszieren. Mit den fluoreszierenden Kataboliten steht eine neue molekulare In-vivo-Sonde für Seneszenzprozesse in Pflanzen zur Verfügung.

Exposure of the filamentous turf green alga Urospora penicilliformis to ambient and artificial ultraviolet radiation (UVR) revealed a considerable resilient species. This explains the ability of this alga to thrive in the middle-upper intertidal zones of the Arctic sea where it is periodically exposed to environmental extremes. A transient UVR effect on photosynthesis under photosynthetically active radiation (PAR) + UV-A and PAR + UV-A + UV-B was found, but dynamic recovery of photoinhibition was observed immediately after reduction of the photon fluence rate of PAR in the absence or presence of background UVR under laboratory and natural solar radiation, respectively. Chlorophylls, carotenoids, and xanthophyll cycle pigments (violaxanthin, antheraxanthin, and zeaxanthin) concentrations were not significantly different between freshly collected samples and filaments exposed to additional laboratory radiation treatment. The ultrastructure of the U. penicilliformis gametophytes showed that the cells are well adapted to UVR. No significant ultrastructural alterations were observed in filaments exposed to different spectral irradiance in the laboratory compared to in situ acclimated specimen. The antioxidant alpha-tocopherol was detected in minute quantity while the search for flavonoid-like compounds was negative. Other UV screening strategies or certain genetically fixed physiological protective mechanism could be operating in this species responsible for their occurrence in higher shoreline and ecological success. Further molecular and biochemical studies are needed to elucidate the stress resistance in this turf alga. There is an indication that the extremely thick cell wall of U. penicilliformis gametophytes covered with mucilage sheath and dense layer of mineral depositions may provide a shield against unfavorable environmental conditions in general and against UVR in particular.

To prevent carbon (C) loss to consumers, trees need to defend their primary production. The tree-internal conflict in resource allocation between growth and defence demands has been the subject of various hypotheses but still requires quantification. A conceptual approach to approximating the C amount dispensable in favour of primary production at the expense of defence is demonstrated which is based on nine defence-related metabolite groups. Quantification is exemplified at the level of sun and shade leaves of adult Fagus sylvatica and Picea abies trees, two species contrasting in foliage type, under oxidative stress as induced by ozone exposure. The difference between maximum and minimum metabolite levels sampled several times throughout four consecutive growing seasons were conceived as dispensable between growth and defence-related metabolism and expressed in proportion of the mean annual gross primary production (GPP) of the foliage. In both species, this proportion amounted to between 2 and 5% of GPP (on a molar C basis). Remaining uncertainties are discussed as concerning functional overlap of substances between growth and defence-related metabolism, estimated classification of metabolite turnover rates and “third-party” trade-offs across C demands. Given the complexity of plant defence, simplification is needed for modelling allocation trade-offs in plants. The presented conceptual approach meets this need in approximating C transfer capacities between competing physiological demands and in stimulating empirical assessments towards mechanistic understanding.

Mesotaenium berggrenii belongs to the few algae that have adapted to live on bare glacial surfaces. Although it is regarded as a cosmopolitan in alpine and polar regions, reports of occurrence are quite sparse. This is probably because of the inconspicuousness of their growing sites, where even mass accumulations only cause a greyish colouration of the white substrate. We characterise a community in the European Alps, including ecophysiological habitat parameters. The field samples consist of two size classes; the smaller one was recognised as M. berggrenii var. alaskana, which contains only one chloroplast per cell after cell division instead of two. This is the first report on the rediscovery of this variety since its description in 1942 and also the first work on its ultrastructure. The results show that M. berggrenii is able to persist in its harsh habitat without formation of cysts, which is in contrast to many snow algal species. The vegetative cells of this cold-adapted species contain high amounts of intraplastidal starch, cytoplasmic lipid bodies as well as many peripheral vacuoles. Organelles, like the Golgi stacks or chloroplasts, are well developed and do not show any signs of morphological reduction, as in cysts. The ecological function of a brownish secondary pigment, a putative polyphenol occurring in high concentrations in vacuoles, is discussed. Screenings by high-performance liquid chromatography reveal the presence of primary photosynthetic pigments characteristic for green algae, but no flavonoid-like phenolic compounds are found.

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Arctic and alpine plants like Oxyria digyna have to face enhanced environmental stress. This study compared leaves from Oxyria digyna collected in the Arctic at Svalbard (78 degrees N) and in the Austrian Alps (47 degrees N) at cellular, subcellular, and ultrastructural levels. Oxyria digyna plants collected in Svalbard had significantly thicker leaves than the samples collected in the Austrian Alps. This difference was generated by increased thickness of the palisade and spongy mesophyll layers in the arctic plants, while epidermal cells had no significant size differences between the two habitats. A characteristic feature of arctic, alpine, and cultivated samples was the occurrence of broad stroma-filled chloroplast protrusions, 2 - 5 microm broad and up to 5 microm long. Chloroplast protrusions were in close spatial contact with other organelles including mitochondria and microbodies. Mitochondria were also present in invaginations of the chloroplasts. A dense network of cortical microtubules found in the mesophyll cells suggested a potential role for microtubules in the formation and function of chloroplast protrusions. No direct interactions between microtubules and chloroplasts, however, were observed and disruption of the microtubule arrays with the anti-microtubule agent oryzalin at 5 - 10 microM did not alter the appearance or dynamics of chloroplast protrusions. These observations suggest that, in contrast to studies on stromule formation in Nicotiana, microtubules are not involved in the formation and morphology of chloroplast protrusions in Oxyria digyna. The actin microfilament-disrupting drug latrunculin B (5 - 10 microM for 2 h) arrested cytoplasmic streaming and altered the cytoplasmic integrity of mesophyll cells. However, at the ultrastructural level, stroma-containing, thylakoid-free areas were still visible, mostly at the concave sides of the chloroplasts. As chloroplast protrusions were frequently found to be mitochondria-associated in Oxyria digyna, a role in metabolite exchange is possible, which may contribute to an adaptation to alpine and arctic conditions.

In leaf mesophyll cells of transgenic Arabidopsis thaliana plants expressing GFP in the chloroplast, stromules (stroma-filled tubules) with a length of up to 20 mu m and a diameter of about 400-600 nm are observed in cells with spaces between the chloroplasts. They appear extremely dynamic, occasionally branched or polymorphic. In order to investigate the effect of temperature on chloroplasts, we have constructed a special temperature-controlled chamber for usage with a light microscope (LM-TCC). This LM-TCC enables presetting of the temperature for investigation directly at the microscope stage with an accuracy of +/- 0.1 degrees C in a temperature range of 0 degrees C to +60 degrees C. With the LM-TCC a temperature-dependent appearance of chloroplast protrusions has been found. These structures have a considerably smaller length-to-diameter ratio than typical stromules and reach a length of 3-5 mu m. At 5-15 degrees C (low temperatures), almost no chloroplast protrusions are observed, but they appear with increasing temperatures. At 35-45 degrees C (high temperatures), numerous chloroplast protrusions with a beaklike appearance extend from a single chloroplast. Interaction of stromules with other organelles has also been investigated by transmission electron microscopy. At 20 degrees C, transverse sections of stromules are frequently observed with a diameter of about 450 nm. A close membrane-to-membrane contact of stromules with the nucleus and mitochondria has been visualised. Golgi stacks and microbodies are found in the spatial vicinity of stromules. At 5 degrees C, virtually no chloroplast protrusions or stromules are observed. At 35 degrees C, chloroplast protrusions are present as broader thylakoid-free stroma-filled areas, resulting in an irregular chloroplast appearance.

The cytology of leaf cells from five different high-alpine plants was studied and compared with structures in chloroplasts from the typical high-alpine plant Ranunculus glacialis previously described as having frequent envelope plus stroma protrusions. The plants under investigation ranged from subalpine/alpine Geum montanum through alpine Geum reptans, Poa alpina var. vivipara, and Oxyria digyna to nival Cerastium uniflorum and R. glacialis. The general leaf structure (by light microscopy) and leaf mesophyll cell ultrastructure (by transmission electron microscopy [TEM]) did not show any specialized structures unique to these mountain species. However, chloroplast protrusion formation could be found in G. reptans and, to a greater extent, in O. digyna. The other species exhibited only a low percentage of such chloroplast structural changes. Occurrence of protrusions in samples of G. montanum and O. digyna growing in a mild climate at about 50 m above sea level was drastically reduced. Serial TEM sections of O. digyna cells showed that the protrusions can appear as rather broad and long appendices of plastids, often forming pocketlike structures where mitochondria and microbodies are in close vicinity to the plastid and to each other. It is suggested that some high-alpine plants may form such protrusions to facilitate fast exchange of molecules between cytoplasm and plastid as an adaptation to the short, often unfavorable vegetation period in the Alps, while other species may have developed different types of adaptation that are not expressed in ultrastructural changes of the plastids.