Psychrophiles or cryophiles (adj. psychrophilic or cryophilic) are extremophilic organisms that are capable of growth and reproduction in low temperatures, ranging from to . 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 mesophiles at intermediate temperatures. Psychrophile is Greek for 'cold-loving', .
Many such organisms are bacteria or archaea, but some eukaryotes such as lichens, snow algae, phytoplankton, 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, 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 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, carotenoids 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 sp., Psychrobacter sp. and members of the genera Halomonas, Pseudomonas, Hyphomonas, and Sphingomonas. Another example is Chryseobacterium greenlandensis, a psychrophile that was found in 120,000-year-old ice.
Umbilicaria antarctica and Xanthoria elegans 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.
Microalgae that live in snow and ice include green, brown, and red algae. Snow algae species such as Chloromonas sp., Chlamydomonas sp., and Chlorella sp. are found in polar environments.