Psychrophiles or cryophiles (adj. ''psychrophilic'' or ''cryophilic'') are

extremophilic An extremophile (from Latin ' meaning "extreme" and Greek ' () meaning "love") is an organism that is able to live (or in some cases thrive) in extreme environments, i.e. environments that make survival challenging such as due to extreme temper ...

organisms that are capable of

growth Growth may refer to: Biology * Auxology, the study of all aspects of human physical growth * Bacterial growth * Cell growth * Growth hormone, a peptide hormone that stimulates growth * Human development (biology) * Plant growth * Secondary growth ...

and

reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual or ...

in low temperatures, ranging from to . They have an optimal growth temperature at . They are found in places that are permanently cold, such as the polar regions and the deep sea. They can be contrasted with thermophiles, which are organisms that thrive at unusually high temperatures, and

mesophile A mesophile is an organism that grows best in moderate temperature, neither too hot nor too cold, with an optimum growth range from . The optimum growth temperature for these organisms is 37°C. The term is mainly applied to microorganisms. Organi ...

s at intermediate temperatures. Psychrophile is Greek for 'cold-loving', . Many such organisms are bacteria or

archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...

, but some

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s such as

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.snow algae,

phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...

, fungi, and wingless midges, are also classified as psychrophiles.

Biology

Habitat

The cold environments that psychrophiles inhabit are ubiquitous on Earth, as a large fraction of the planetary surface experiences temperatures lower than 10 °C. They are present in

permafrost Permafrost is ground that continuously remains below 0 °C (32 °F) for two or more years, located on land or under the ocean. Most common in the Northern Hemisphere, around 15% of the Northern Hemisphere or 11% of the global surface ...

, polar ice, glaciers, snowfields and deep ocean waters. These organisms can also be found in pockets of sea ice with high salinity content. Microbial activity has been measured in soils frozen below −39 °C. In addition to their temperature limit, psychrophiles must also adapt to other extreme environmental constraints that may arise as a result of their habitat. These constraints include high pressure in the deep sea, and high salt concentration on some sea ice.

Adaptations

Psychrophiles are protected from freezing and the expansion of ice by ice-induced

desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...

and vitrification (glass transition), as long as they cool slowly. Free living cells desiccate and vitrify between −10 °C and −26 °C. Cells of multicellular organisms may vitrify at temperatures below −50 °C. The cells may continue to have some metabolic activity in the extracellular fluid down to these temperatures, and they remain viable once restored to normal temperatures. They must also overcome the stiffening of their lipid cell membrane, as this is important for the survival and functionality of these organisms. To accomplish this, psychrophiles adapt lipid membrane structures that have a high content of short, unsaturated fatty acids. Compared to longer saturated fatty acids, incorporating this type of fatty acid allows for the lipid cell membrane to have a lower melting point, which increases the fluidity of the membranes. In addition,

carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic compound, organic pigments that are produced by plants and algae, as well as several bacteria, and Fungus, fungi. Carotenoids give the characteristic color to pumpki ...

s are present in the membrane, which help modulate the fluidity of it. Antifreeze proteins are also synthesized to keep psychrophiles' internal space liquid, and to protect their DNA when temperatures drop below water's freezing point. By doing so, the protein prevents any ice formation or recrystallization process from occurring. The enzymes of these organisms have been hypothesized to engage in an activity-stability-flexibility relationship as a method for adapting to the cold; the flexibility of their enzyme structure will increase as a way to compensate for the freezing effect of their environment. Certain cryophiles, such as Gram-negative bacteria ''Vibrio'' and ''Aeromonas'' spp., can transition into a viable but nonculturable (VBNC) state. During VBNC, a micro-organism can respire and use substrates for metabolism – however, it cannot replicate. An advantage of this state is that it is highly reversible. It has been debated whether VBNC is an active survival strategy or if eventually the organism's cells will no longer be able to be revived. There is proof however it may be very effective – Gram positive bacteria Actinobacteria have been shown to have lived about 500,000 years in the permafrost conditions of Antarctica, Canada, and Siberia.

Taxonomic range

Psychrophiles include bacteria, lichens, snow algae, phytoplankton, fungi, and insects. Among the bacteria that can tolerate extreme cold are ''

Arthrobacter ''Arthrobacter'' (from the Greek, "jointed small stick”) is a genus of bacteria that is commonly found in soil. All species in this genus are Gram-positive obligate aerobes that are rods during exponential growth and cocci in their stationary ...

'' sp., ''

Psychrobacter ''Psychrobacter'' is a genus of Gram-negative, osmotolerant, oxidase-positive, psychrophilic or psychrotolerant, aerobic bacteria which belong to the family Moraxellaceae and the class Gammaproteobacteria. The shape is typically cocci or cocco ...

'' sp. and members of the genera ''

Halomonas ''Halomonas'' is a genus of halophilic (salt-tolerating) bacteria. It grows over the range of 5 to 25% NaCl. The type species of this genus is '' Halomonas elongata''. Description Members of ''Halomonas'' are Gram-negative, rod-shaped bacteria, ...

'', '' Pseudomonas'', ''

Hyphomonas In taxonomy, ''Hyphomonas'' is a genus of the Hyphomonadaceae.See the NCBI The National Center for Biotechnology Information (NCBI) is part of the United States National Library of Medicine (NLM), a branch of the National Institutes of Healt ...

'', and '' Sphingomonas''. Another example is ''

Chryseobacterium greenlandensis ''Chryseobacterium aquaticum'' is a Gram-negative, non- spore-forming and non-motile bacteria Non-motile bacteria are bacteria species that lack the ability and structures that would allow them to propel themselves, under their own power, thr ...

'', a psychrophile that was found in 120,000-year-old ice. ''

Umbilicaria antarctica Rock tripe is the common name for various lichens of the genus ''Umbilicaria'' that grow on rocks. They are widely distributed, including on bare rock in Antarctica, and throughout northern parts of North America such as New England and the Rocky ...

'' and ''

Xanthoria elegans ''Xanthoria elegans'', commonly known as the elegant sunburst lichen, is a lichenized species of fungus in the genus ''Xanthoria'', family Teloschistaceae. Recognized by its bright orange or red pigmentation, this species grows on rocks, often ne ...

'' are lichens that have been recorded photosynthesizing at temperatures ranging down to −24 °C, and they can grow down to around −10 °C. Some multicellular eukaryotes can also be metabolically active at sub-zero temperatures, such as some conifers; those in the '' Chironomidae'' family are still active at −16 °C. Chlamydomonas3 (Antarctique)

Microalgae Microalgae or microphytes are microscopic algae invisible to the naked eye. They are phytoplankton typically found in freshwater and marine systems, living in both the water column and sediment. They are unicellular species which exist indiv ...

that live in snow and ice include green, brown, and red algae. Snow algae species such as ''

Chloromonas ''Chloromonas'' is a genus of green algae in the family Chlamydomonadaceae. It is closely related to the model green algae, ''Chlamydomonas'', and distinguished mainly through the absence of a pyrenoid Pyrenoids are sub-cellular micro-compart ...

sp.'', ''

Chlamydomonas ''Chlamydomonas'' is a genus of green algae consisting of about 150 speciesSmith, G.M. 1955 ''Cryptogamic Botany Volume 1. Algae and Fungi'' McGraw-Hill Book Company Inc of unicellular flagellates, found in stagnant water and on damp soil, ...

sp.'', and '' Chlorella sp.'' are found in polar environments. Some

can tolerate extremely cold temperatures and high salinities that occur in brine channels when

sea ice Sea ice arises as seawater freezes. Because ice is less dense than water, it floats on the ocean's surface (as does fresh water ice, which has an even lower density). Sea ice covers about 7% of the Earth's surface and about 12% of the world's oce ...

forms in polar oceans. Some examples are diatoms like ''

Fragilariopsis cylindrus ''Fragilariopsis cylindrus'' is a Pennales, pennate sea-ice diatom that is found native in the Argentine Sea and Antarctic waters, with a pH of 8.1-8.4. It is regarded as an Bioindicator, indicator species for polar water. Description ''Fragil ...

'', ''Nitzchia lecointeii'', ''Entomoneis kjellmanii'', ''Nitzchia stellata'', ''Thalassiosira australis'', ''Berkelaya adeliense'', and ''Navicula glaciei''. '' Penicillium'' is a genus of fungi found in a wide range of environments including extreme cold. Among the psychrophile insects, the

Grylloblattidae Grylloblattidae, commonly known as the icebugs, or ice crawlers, is a family of extremophile (psychrophile) and wingless insects that live in the cold on top of mountains and the edges of glaciers. They belong, along with Mantophasmatidae (rock c ...

or ice crawlers, found on mountaintops, have optimal temperatures between 1-4 °C. The wingless midge (Chironomidae) '' Belgica antarctica'' can tolerate salt, being frozen and strong ultraviolet, and has the smallest known genome of any insect. The small genome, of 99 million

base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...

s, is thought to be adaptive to extreme environments.

Psychrotrophic bacteria

Psychrotrophic microbes are able to grow at temperatures below , but have better growth rates at higher temperatures. Psychrotrophic bacteria and fungi are able to grow at refrigeration temperatures, and can be responsible for food spoilage and as foodborne pathogens such as '' Yersinia''. They provide an estimation of the product's shelf life, but also they can be found in soils, in surface and deep sea waters, in Antarctic ecosystems, and in foods. Psychrotrophic bacteria are of particular concern to the dairy industry. Most are killed by

pasteurization Pasteurization or pasteurisation is a process of food preservation in which packaged and non-packaged foods (such as milk and fruit juices) are treated with mild heat, usually to less than , to eliminate pathogens and extend shelf life. The ...

; however, they can be present in milk as post-pasteurization contaminants due to less than adequate sanitation practices. According to the Food Science Department at Cornell University, psychrotrophs are bacteria capable of growth at temperatures at or less than . At freezing temperatures, growth of psychrotrophic bacteria becomes negligible or virtually stops. All three subunits of the RecBCD enzyme are essential for physiological activities of the enzyme in the Antarctic '' Pseudomonas syringae'', namely, repairing of DNA damage and supporting the growth at low temperature. The RecBCD enzymes are exchangeable between the psychrophilic ''P. syringae'' and the mesophilic ''E. coli'' when provided with the entire protein complex from same species. However, the RecBC proteins (RecBCPs and RecBCEc) of the two bacteria are not equivalent; the RecBCEc is proficient in DNA recombination and repair, and supports the growth of ''P. syringae'' at low temperature, while RecBCPs is insufficient for these functions. Finally, both helicase and nuclease activity of the RecBCDPs are although important for DNA repair and growth of ''P. syringae'' at low temperature, the RecB-nuclease activity is not essential in vivo.

Psychrophilic microalgae

Broken pack ice with cryopelagic antarctic diatoms

Microscopic algae that can tolerate extremely cold temperatures can survive in snow, ice, and very cold seawater. On snow, cold-tolerant algae can bloom on the snow surface covering land, glaciers, or sea ice when there is sufficient light. These snow algae darken the surface of the snow and can contribute to snow melt. In seawater, phytoplankton that can tolerate both very high salinities and very cold temperatures are able to live in sea ice. One example of a psychrophilic phytoplankton species is the ice-associated diatom ‘’

’’. Phytoplankton living in the cold ocean waters near Antarctica often have very high protein content, containing some of the highest concentrations ever measured of enzymes like Rubisco.

Psychrotrophic insects

Insects that are psychrotrophic can survive cold temperatures through several general mechanisms (unlike opportunistic and chill susceptible insects): (1) chill tolerance, (2) freeze avoidance, and (3) freeze tolerance. Chill tolerant insects succumb to freezing temperatures after prolonged exposure to mild or moderate freezing temperatures. Freeze avoiding insects can survive extended periods of time at sub-freezing temperatures in a supercooled state, but die at their supercooling point. Freeze tolerant insects can survive ice crystal formation within their body at sub-freezing temperatures. Freeze tolerance within insects is argued to be on a continuum, with some insect species exhibiting partial (e.g., '' Tipula paludosa'', ''

Hemideina thoracica ''Hemideina thoracica'', commonly known as the Auckland tree wētā or tokoriro is a cricket-like insect (within the family Anostostomatidae). It is endemic to New Zealand and is found over most of the North Island, except for the Wellington re ...

'' ), moderate (e.g., '' Cryptocercus punctulatus''), and strong freezing tolerance (e.g., ''

Eurosta solidaginis The goldenrod gall fly (''Eurosta solidaginis''), also known as the goldenrod ball gallmaker, is a species of fly native to North America. The species is best known for the characteristic galls it forms on several species in the ''Solidago'', or ...

'' and ''

Syrphus ribesii ''Syrphus ribesii'' is a very common Holarctic species of hoverfly. Its larvae feed on aphids. In common with many other species of hoverfly, males have the eyes meeting on the top of the head, whilst females have their eyes widely separated. De ...

'''')'', and other insect species exhibiting freezing tolerance with low supercooling point (e.g., ''

Pytho deplanatus Pytho may refer to: * Delphi or Pytho, a sacred precinct that served as the seat of the oracle Pythia * ''Pytho'' (beetle), a genus of dead-log beetles * Pytho, an order of devils in the classification of demons See also * Python (mythology) ...

'').

Psychrophile versus psychrotroph

In 1940, ZoBell and Conn stated that they had never encountered "true psychrophiles" or organisms that grow best at relatively low temperatures. In 1958, J. L. Ingraham supported this by concluding that there are very few or possibly no bacteria that fit the textbook definitions of psychrophiles. Richard Y. Morita emphasizes this by using the term ''psychrotroph'' to describe organisms that do not meet the definition of psychrophiles. The confusion between the terms ''psychrotrophs'' and ''psychrophiles'' was started because investigators were unaware of the thermolability of psychrophilic organisms at the laboratory temperatures. Due to this, early investigators did not determine the cardinal temperatures for their isolates. The similarity between these two is that they are both capable of growing at zero, but optimum and upper temperature limits for the growth are lower for psychrophiles compared to psychrotrophs. Psychrophiles are also more often isolated from permanently cold habitats compared to psychrotrophs. Although psychrophilic enzymes remain under-used because the cost of production and processing at low temperatures is higher than for the commercial enzymes that are presently in use, the attention and resurgence of research interest in psychrophiles and psychrotrophs will be a contributor to the betterment of the environment and the desire to conserve energy.