Psychrophilic enzymes: hot topics in cold adaptation

More than three-quarters of the Earth's surface is occupied by cold ecosystems, including the ocean depths, and polar and alpine regions. These permanently cold environments have been successfully colonized by a class of extremophilic microorganisms that are known as psychrophiles (which literally means cold-loving). The ability to thrive at temperatures that are close to, or below, the freezing point of water requires a vast array of adaptations to maintain the metabolic rates and sustained growth compatible with life in these severe environmental conditions.     

Genomic convergence underlying high-altitude adaptation in alpine plants

Evolutionary convergence is one of the most striking examples of adaptation driven by natural selection.However, genomic evidence for convergent adaptation to extreme environments remains scarce.Here, we assembled reference genomes of two alpine plants, Saussurea obvallata(Asteraceae)and Rheum alexandrae(Polygonaceae), with 37,938 and 61,463 annotated protein-coding genes. By integrating an additional five alpine genomes,we elucidated genomic convergence underlying high-altitude adaptation in al...     

ADAPTATIONS OF TERRESTRIAL ARTHROPODS TO THE ALPINE ENVIRONMENT

1. The climate changes drastically above the timberline. Diverse adaptations have been evolved by insects and other terrestrial arthropods to survive the alpine environment. The fitness of each species depends on a combination of different factors in accordance with their special habitats. 2. Morphological adaptations such as reduced body-size, are known from a number of alpine insects, increasing their possibility to find sheltered microhabitats. Selection for reduced body size in Andean Phulia spp. butterflies is probably a result of their rigorous environment. Wing atrophy, which is also known in insects from other extreme environments, is widespread in alpine species. In several terrestrial arthropods the absorption of solar radiation is increased by melanism. Increased pubescence, protecting against the loss of heat, is known in alpine butterflies and bumblebees. 3. Several behavioural adaptations are described. Thermoregulatory behaviour is important in many species to raise their body temperatures. Alpine butterflies orient the dark basis of their wings perpendicular to the rays of the sun. Body temperatures of 30 °C may be required for flight. To increase their activities many alpine terrestrial arthropods seek warmer microhabitats in the vegetation and under rocks. The adaptive advantage of nocturnal activity as observed in several species, may be to maintain the water balance or to avoid predation. 4. Tropical alpine terrestrial arthropods are faced with special problems. The large diel temperature fluctuations require cold-hardiness during the night and tolerance to heat during the day. Many species seek sheltered microhabitats under rocks and in vegetation. 5. Due to low precipitation and high evaporation rates many mountain areas are extremely dry. High resistance to desiccation may be very important to alpine species, and in particular to tropical species. Rates of water loss at low relative humidities are comparable to those of desert arthropods. 6. As an adaptation to the cold alpine summers several species of terrestrial arthropods require more than one year to complete their life-cycles. Special to these species is their adaptation to low temperatures in two or more overwintering stages. In spite of their cold surroundings several species have univoltine life cycles, frequently combined with highly specialized adaptations. Increased metabolic rates as a compensation to low temperatures may be widespread in alpine species, but few data are available. 7. Cold tolerance is of particular importance in temperature alpine species. Winter survival in Collembola and Acari depends on supercooling. Great seasonal variations have been observed in a number of species. Freezing tolerance is also known from alpine insects, e.g. in some species of beetles. At high latitudes alpine species must endure periods of up to eight or nine month at low temperatures during hibernation. Anaerobiosis is known from species that are enclosed in ice, with lactate as the main end product of metabolism.     

Alpine microorganisms: useful tools for low-temperature bioremediation

Cold environments, including polar and alpine regions, are colonized by a wide diversity of microorganisms able to thrive at low temperatures. There is evidence of a wide range of metabolic activities in alpine cold ecosystems. Like polar microorganisms, alpine microorganisms play a key ecological role in their natural habitats for nutrient cycling, litter degradation, and many other processes. A number of studies have demonstrated the capacity of alpine microorganisms to degrade efficiently a wide range of hydrocarbons, including phenol, phenol-related compounds and petroleum hydrocarbons, and the feasibility of low-temperature bioremediation of European alpine soils by stimulating the degradation capacity of indigenous microorganisms has also been shown.     

低温微生物及其酶类的研究概况

广泛分布在地球寒冷生境 ,如南北两极、高山、深海以及冰川中的低温微生物 ,不但为研究低温生态系统、生命起源与进化以及生物适冷机制提供了丰富的材料 ,同时在生物工程方面也具有潜在的巨大开发价值。国内外越来越多的科研人员对低温微生物及其产物的研究表现出了浓厚的兴趣。关于细胞膜和低温酶的研究 ,是目前微生物适冷机制研究中的 2个热点。就低温微生物的研究现状和适冷机制以及低温酶类的研究进行了综述。     

Adaptation of fungi,including yeasts,to cold environments

A wide range of cold environments exist, with an equally broad variety of fungi and yeasts that have adapted to such environments. These adaptations, which affect membranes, enzymes and other cellular components, such as radical scavenging molecules, display a great potential for exploitation in biotechnology. Alterations have been detected in membrane lipids, with an increase in fatty acid unsaturated bonds that enhance their fluidity. We report new data on the different phospholipid composition in membrane lipids in the same fungal species from both Antarctic and temperate regions. The decrease in temperature causes intracellular oxidative stress by inducing the generation of reactive oxygen species. We report the results of the first analysis of the non-enzymatic antioxidant response and phenolic compound production by an Antarctic strain of Geomyces pannorum. A survey on yeasts from the cryosphere is reported with a focus on their adaptation to a cold environment. Some studies have shown that the number of macrofungi in glacier forefronts rises as deglaciation increases. The survival success of many plants in such areas may be attributed to their mycorrhizal associations. We highlighted the macrofungal biodiversity of some Italian alpine habitats, in which we Inocybe microfastigiata, Laccaria montana and Lactarius salicis-herbaceae were recorded for the first time in Lombardy (Italy).     

Psycrophilic organisms in snow and ice]

Psychrophilic and psycrotrophic organisms are important in global ecology as a large proportion of our planet is cold. Two-third of sea-water covering more than 70% of Earth is cold deep sea water with temperature around 2 degrees C, and more than 90% of freshwater is in polar ice-sheets and mountain glaciers. Though biological activity in snow and ice had been believed to be extremely limited, various specialized biotic communities were recently discovered at glaciers of various part of the world. The glacier is relatively simple and closed ecosystem with special biotic community containing various psychrophilic and psycrotrophic organisms. Since psychrophilic organisms was discovered in the deep ice-core recovered from the antarctic ice-sheet and a lake beneath it, snow and ice environments in Mars and Europa are attracting a great deal of scientific attention as possible extraterrestrial habitats of life. This paper briefly reviews the results of the studies on ecology of psychrophilic organisms living in snow and ice environments and their physiological and biochemical adaptation to low temperature.     

Ecophysiology of Antarctic vascular plants

Most of the ice and snow-free land in the Antarctic summer is found along the Antarctic Peninsula and adjacent islands and coastal areas of the continent. This is the area where most of the Antarctic vegetation is found. Mean air temperature tends to be above zero during the summer in parts of the Maritime Antarctic. The most commonly found photosynthetic organisms in the Maritime Antarctic and continental edge are lichens (around 380 species) and bryophytes (130 species). Only two vascular plants, Deschampsia antarctica Desv. and Colobanthus quitensis (Kunth) Bartl., have been able to colonize some of the coastal areas. This low species diversity, compared with the Arctic, may be due to permanent low temperature and isolation from continental sources of propagules. The existence of these plants in such a permanent harsh environment makes them of particular interest for the study of adaptations to cold environments and mechanisms of cold resistance in plants. Among these adaptations are high freezing resistance, high resistance to light stress and high photosynthetic capacity at low temperature. In this paper, the ecophysiology of the two vascular plants is reviewed, including habitat characteristics, photosynthetic properties, cold resistance, and biochemical adaptations to cold.     

Vertical and seasonal variation of phytoplankton photosynthesis in a brown-water lake with winter ice cover

SUMMARY. Unlike previously studied lakes with prolonged winter ice and snow cover, Lake Paajarvi, southern Finland, has a high humus content and consequently differs in both the quantity and quality of light penetration into its waters. Moreover, the range of temperature fluctuation and the degree of development of thermal stratification are greater in Paajarvi, and this increased environmental heterogeneity apparently stimulates diversity in the phytoplankton community, especially in the seasonal succession of species. Differences in the photosynthetic capacity of algae from different depths in the water column were not great. This is attributed to the extremely shallow euphotic zone, algae circulating freely through the steep light gradient and sedimenting rapidly once they pass through the thermocline into the hypolimnion. It is suggested that 'adaptation' of phytoplankton to the great seasonal changes in irradiance is achieved largely by successive growths of different species in the community, and that the adaptations and vertical migrations by individual algal species, which have been reported from polar and high alpine lakes, may be of secondary importance in Pääjärvi. The species successions in Pääjärvi produce changes in the pigment content of algae similar to those reported from polar and high alpine lakes, confirming that change in pigmentation is an important mechanism in light adaptation, whether at community or individual level. Algal pigment content was particularly high at the end of the long period of winter ice cover, indicating a degree of adaptation to the prolonged low-light conditions, which produced the extremely high photosynthetic capacities measured at this time. However, phytoplankton production at any irradiance was primarily determined by biomass.     

四裂红景天与长鳞红景天叶片中抗氧化系统随海拔梯度的变化

以分布于乌鲁木齐河源区天山中段不同海拔高度的四裂红景天 (Rhodiolaquadrifida) (35 0 0~ 380 5m) 及长鳞红景天 (R.gelida) (380 5~ 4 0 10m) 为试材, 通过对比分析两种红景天在一连续的海拔梯度上种内及种间叶片膜脂过氧化及抗氧化保护系统的变化, 初步探讨植物对于高山极端环境的适应机制。结果显示, 分布于海拔较高的长鳞红景天其叶片中膜脂过氧化产物丙二醛 (MDA) 的含量明显高于分布海拔较低的四裂红景天, 而两种红景天种内膜脂过氧化水平则没有随海拔升高呈现出明显差异。此外, 抗氧化保护酶CAT, POD, SOD, GR和ASA_POD 的活性与非酶促抗氧化剂ASA、GSH的含量不仅在种间存在有明显差异, 长鳞红景天中明显高于四裂红景天, 而且在两种红景天种内亦随海拔升高有不同程度地提高。表明当海拔升高时, 虽然环境条件渐趋恶劣, 对植物造成的氧化胁迫增强, 但红景天脂膜保护系统的功能亦相应加强, 从而增强了其抵抗逆境胁迫的能力。其中以过氧化物酶POD的活性变化最为显著 :在 380 5m以下海拔区, 于四裂红景天叶片中均未检测到该酶活性, 而分布海拔较高的长鳞红景天叶片中该酶活性则随海拔升高明显增强, 推测POD可能在红景天适应特殊生境中起着重要的作用, 亦可能与高海拔区长鳞红景天取代四裂红景天有关。     

How do vascular plants perform photosynthesis in extreme environments? An integrative ecophysiological and biochemical story

In this work, we review the physiological and molecular mechanisms that allow vascular plants to perform photosynthesis in extreme environments, such as deserts, polar and alpine ecosystems. Specifically, we discuss the morpho/anatomical, photochemical and metabolic adaptive processes that enable a positive carbon balance in photosynthetic tissues under extreme temperatures and/or severe water‐limiting conditions in C₃ species. Nevertheless, only a few studies have described the in situ functioning of photoprotection in plants from extreme environments, given the intrinsic difficulties of fieldwork in remote places. However, they cover a substantial geographical and functional range, which allowed us to describe some general trends. In general, photoprotection relies on the same mechanisms as those operating in the remaining plant species, ranging from enhanced morphological photoprotection to increased scavenging of oxidative products such as reactive oxygen species. Much less information is available about the main physiological and biochemical drivers of photosynthesis: stomatal conductance (g_s), mesophyll conductance (g_m) and carbon fixation, mostly driven by RuBisCO carboxylation. Extreme environments shape adaptations in structures, such as cell wall and membrane composition, the concentration and activation state of Calvin–Benson cycle enzymes, and RuBisCO evolution, optimizing kinetic traits to ensure functionality. Altogether, these species display a combination of rearrangements, from the whole‐plant level to the molecular scale, to sustain a positive carbon balance in some of the most hostile environments on Earth.     

Metabolic coping strategies in cold tolerant reptiles

The reality of metabolic cold adaptation in ectotherms is disputed. Reptiles differ in many important aspects of their thermal ecology from aquatic ectotherms, and their thermophysiology is therefore difficult to analyze in conventional terms. Evidence is presented in support of the view that both phenotypic metabolic cold acclimation and genotypic metabolic cold adaptation are important coping strategies in cold tolerant reptiles. Functional explanations of these paradoxical adaptations are discussed in the light of a multidimensional energetic model of the reptilian thermal niche.     

Psychrophiles and polar regions

Most reviews of microbial life in cold environments begin with a lament of how little is known about the psychrophilic (cold-loving) inhabitants or their specific adaptations to the cold. This situation is changing, as research becomes better focused by new molecular genetic (and other) approaches, by awareness of accelerated environmental change in polar regions, and by strong interest in the habitability of frozen environments elsewhere in the solar system. This review highlights recent discoveries in molecular adaptation, biodiversity and microbial dynamics in the cold, along with the concept of eutectophiles, organisms living at the critical interface inherent to the phase change of water to ice.     

宽果苁菔适应昼夜变化ESTs的分离与分析

以流石滩地区植物广布种宽果苁菔为实验材料,应用磁珠介导的抑制消减杂交方法,分离昼夜差异基因。随机挑选了136个差异ESTs克隆并进行测序,测序结果使用Blast2go程序进行功能注释和分析。结果表明绝大部分ESTs功能与稳定细胞状态和抗性响应相关,其次与物质能量代谢和信号传导相关,宽果苁菔适应昼夜变化过程的机制非常复杂,本研究为进一步研究高山植物适应流石滩恶劣环境的机制奠定了基础。     

Carbon acquisition and water relations of lichens in polar regions — potentials and limitations

Primary production in lichens, as in all plants, is the result of gains and losses. Losses of carbon may be high under extreme environmental conditions. In the harsh climate of polar regions lichens use much carbon for maintenance and in their stress response. Thallus biomass is preserved through periods of darkness, extremely low temperatures and snow cover. On the other hand, lichens are photosynthetically active at temperatures below 0°C. They can make use of melted water from snow and ice and take up water vapour from snow. Physiological adaptations and environmental conditions under which growth is possible are discussed in terms of our recent results. A concept model describes the main parameters that influence carbon acquisition, growth and water relations of lichens in polar regions.     

Components of Pink Snow Mould Resistance in Winter Wheat are Expressed Prior to Cold Hardening and in Detached Leaves

Resistance to pink snow mould, caused by Microdochium nivale, was investigated in four resistant winter wheat lines from the USDA World Cereal Collection (CI9342, CI14106, PI173440 and PI181268) and three Nordic wheat lines (Bjørke, Rida and V1004). Pink snow mould resistance was tested in non‐hardened and cold‐hardened plants incubated under artificial snow cover and in detached leaf segments mounted on water agar and incubated at either 3°C in darkness or at room temperature with light during the day. The wheat lines CI9342, CI14106 and PI181268 were more resistant than the Nordic lines, both before and after cold hardening. Thus, although cold hardening strongly increases the level of snow mould resistance in all the wheat lines, some resistance mechanisms are also present prior to cold hardening in some of the resistant lines. CI9342, CI14106 and PI181268 also had a higher level of resistance than the other lines in the detached leaf assay, indicating that these lines have some resistance mechanisms acting in the leaves. The resistance of PI173440 was expressed only in intact hardened plants and not in non‐hardened plants or in detached leaves. This indicates that this line relies on cold hardening‐related changes in the crown for its resistance. In the detached leaf assay the rate of lesion development varied greatly between leaves of different order. The highest correlation with the whole plant test was obtained when using secondary leaves and incubation at 3°C in the dark.     

Life‐history traits and physiological limits of the alpine fly Drosophila nigrosparsa (Diptera: Drosophilidae): A comparative study

Interspecific variation in life‐history traits and physiological limits can be linked to the environmental conditions species experience, including climatic conditions. As alpine environments are particularly vulnerable under climate change, we focus on the montane‐alpine fly Drosophila nigrosparsa. Here, we characterized some of its life‐history traits and physiological limits and compared these with those of other drosophilids, namely Drosophila hydei, Drosophila melanogaster, and Drosophila obscura. We assayed oviposition rate, longevity, productivity, development time, larval competitiveness, starvation resistance, and heat and cold tolerance. Compared with the other species assayed, D. nigrosparsa is less fecund, relatively long‐living, starvation susceptible, cold adapted, and surprisingly well heat adapted. These life‐history characteristics provide insights into invertebrate adaptations to alpine conditions which may evolve under ongoing climate change.