Haloarchaea


Haloarchaea are a class of the Euryarchaeota, found in water saturated or nearly saturated with salt. Halobacteria are now recognized as archaea, rather than bacteria and are one of the largest groups. The name 'halobacteria' was assigned to this group of organisms before the existence of the domain Archaea was realized, and while valid according to taxonomic rules, should be updated. Halophilic archaea are generally referred to as haloarchaea to distinguish them from halophilic bacteria.
These microorganisms are members of the halophile community, in that they require high salt concentrations to grow, with most species requiring more than 2.0M NaCl for growth and survival. They are a distinct evolutionary branch of the Archaea distinguished by the possession of ether-linked lipids and the absence of murein in their cell walls.
Haloarchaea can grow aerobically or anaerobically. Parts of the membranes of haloarchaea are purplish in color, and large blooms of haloarchaea appear reddish, from the pigment bacteriorhodopsin, related to the retinal pigment rhodopsin, which it uses to transform light energy into chemical energy by a process unrelated to chlorophyll-based photosynthesis. The water of Lonar lake in Maharashtra’s Buldhana district due to the large presence of haloarchaea microbes in the salty water. Further, organisms like zooplankton and crayfish consume these microbes and thereafter serve as carotenoid-rich food sources themselves for larger organisms like flamingos.
Haloarchaea have a potential to solubilize phosphorus. Phosphorus-solubilizing halophilic archaea may well play a role in P nutrition to vegetation growing in hypersaline soils. Haloarchaea may also have applications as inoculants for crops growing in hypersaline regions.

Taxonomy

The extremely halophilic, aerobic members of Archaea are classified within the family Halobacteriaceae, order Halobacteriales in Class III. Halobacteria of the phylum Euryarchaeota. As of May 2016, the family Halobacteriaceae comprises 213 species in 50 genera.
Species list
Domain: Archaea
Euryarchaeota
  • Halobacteria
  • *Halobacteriales
  • **Halobacteriaceae
  • ***Haladaptatus
  • **** Haladaptatus cibarius
  • **** Haladaptatus litoreus
  • **** Haladaptatus pallidirubidus
  • **** Haladaptatus paucihalophilus
  • *** Halalkalicoccus
  • **** Halalkalicoccus jeotgali
  • **** Halalkalicoccus paucihalophilus
  • **** Halalkalicoccus tibetensis
  • *** "Halanaeroarchaeum"
  • **** "Halanaeroarchaeum sulfurireducens"
  • *** Halapricum
  • **** Halapricum salinum
  • *** Halarchaeum
  • **** Halarchaeum acidiphilum
  • **** Halarchaeum grantii
  • **** Halarchaeum nitratireducens
  • **** Halarchaeum rubridurum
  • **** Halarchaeum salinum
  • **** Halarchaeum solikamskense
  • *** Haloarchaeobius
  • **** Haloarchaeobius amylolyticus
  • **** Haloarchaeobius baliensis
  • **** Haloarchaeobius iranensis
  • **** Haloarchaeobius litoreus
  • **** Haloarchaeobius salinus
  • *** Haloarcula
  • **** Haloarcula amylolytica
  • **** Haloarcula argentinensis
  • **** Haloarcula hispanica
  • **** Haloarcula japonica
  • **** Haloarcula marismortui
  • **** Haloarcula quadrata
  • **** Haloarcula salaria
  • **** Haloarcula tradensis
  • **** Haloarcula vallismortis
  • *** Halobacterium
  • ****Halobacterium sp. NRC-1
  • ****Halobacterium jilantaiense
  • ****Halobacterium noricense
  • ****"Halobacterium piscisalsi"
  • ****Halobacterium rubrum
  • ****Halobacterium salinarum
  • *** Halobaculum
  • **** Halobaculum gomorrense
  • **** Halobaculum magnesiiphilum
  • *** Halobellus
  • **** Halobellus clavatus
  • **** Halobellus inordinatus
  • **** Halobellus limi
  • **** Halobellus litoreus
  • **** Halobellus ramosii
  • **** Halobellus rarus
  • **** Halobellus rufus
  • **** Halobellus salinus
  • *** Halobiforma
  • **** Halobiforma haloterrestris
  • **** Halobiforma lacisalsi
  • **** Halobiforma nitratireducens
  • *** Halocalculus
  • **** Halocalculus aciditolerans
  • *** Halococcus
  • **** Halococcus agarilyticus
  • **** Halococcus dombrowskii
  • **** Halococcus hamelinensis
  • **** Halococcus morrhuae
  • **** Halococcus qingdaonensis
  • **** Halococcus saccharolyticus
  • **** Halococcus salifodinae
  • **** Halococcus sediminicola
  • **** Halococcus thailandensis
  • *** Haloferax
  • **** Haloferax alexandrinus
  • **** Haloferax chudinovii
  • **** Haloferax denitrificans
  • **** Haloferax elongans
  • **** Haloferax gibbonsii
  • **** Haloferax larsenii
  • **** Haloferax lucentense
  • **** Haloferax mediterranei
  • **** Haloferax mucosum
  • **** Haloferax namakaokahaiae
  • **** Haloferax prahovense
  • **** Haloferax sulfurifontis
  • **** Haloferax volcanii
  • *** Halogeometricum
  • **** Halogeometricum borinquense
  • **** Halogeometricum limi
  • **** Halogeometricum pallidum
  • **** Halogeometricum rufum
  • *** Halogranum
  • **** Halogranum amylolyticum
  • **** Halogranum gelatinilyticum
  • **** Halogranum rubrum
  • **** Halogranum salarium
  • *** Halohasta
  • **** Halohasta litorea
  • **** Halohasta litchfieldiae
  • *** Halolamina
  • **** Halolamina pelagica
  • **** Halolamina rubra
  • **** Halolamina salifodinae
  • **** Halolamina salina
  • **** Halolamina sediminis
  • *** Halomarina
  • **** Halomarina oriensis
  • *** Halomicroarcula
  • **** Halomicroarcula limicola
  • **** Halomicroarcula pellucida
  • **** Halomicroarcula salina
  • *** Halomicrobium
  • **** Halomicrobium katesii
  • **** Halomicrobium mukohataei
  • **** Halomicrobium zhouii
  • *** Halonotius
  • **** Halonotius pteroides
  • *** "Haloparvum"
  • **** "Haloparvum sedimenti"
  • *** Halopelagius
  • **** Halopelagius inordinatus
  • **** Halopelagius fulvigenes
  • **** Halopelagius longus
  • *** Halopenitus
  • **** Halopenitus malekzadehii
  • **** Halopenitus persicus
  • **** Halopenitus salinus
  • *** Halopiger
  • **** Halopiger aswanensis
  • **** Halopiger salifodinae
  • **** Halopiger xanaduensis
  • *** Haloplanus
  • **** Haloplanus aerogenes
  • **** Haloplanus litoreus
  • **** Haloplanus natans
  • **** Haloplanus ruber
  • **** Haloplanus salinus
  • **** Haloplanus vescus
  • *** Haloquadratum
  • **** Haloquadratum walsbyi
  • *** Halorhabdus
  • **** Halorhabdus tiamatea
  • **** Halorhabdus utahensis
  • **** "Halorhabdus rudnickae"
  • *** Halorientalis
  • **** Halorientalis brevis
  • **** Halorientalis persicus
  • **** Halorientalis regularis
  • *** Halorubellus
  • **** Halorubellus litoreus
  • **** Halorubellus salinus
  • *** Halorubrum
  • **** Halorubrum aquaticum
  • **** Halorubrum aidingense
  • **** Halorubrum alkaliphilum
  • **** Halorubrum arcis
  • **** Halorubrum californiense
  • **** Halorubrum chaoviator
  • **** Halorubrum cibi
  • **** Halorubrum coriense
  • **** Halorubrum distributum
  • **** Halorubrum ejinorense
  • **** Halorubrum ezzemoulense
  • **** Halorubrum gandharaense
  • **** Halorubrum halodurans
  • **** Halorubrum halophilum
  • **** Halorubrum kocurii
  • **** Halorubrum lacusprofundi
  • **** Halorubrum laminariae
  • **** Halorubrum lipolyticum
  • **** Halorubrum litoreum
  • **** Halorubrum luteum
  • **** Halorubrum orientale
  • **** Halorubrum persicum
  • **** Halorubrum rubrum
  • **** "Halorubrum rutilum"
  • **** Halorubrum saccharovorum
  • **** Halorubrum salinum
  • **** Halorubrum sodomense
  • **** Halorubrum tebenquichense
  • **** Halorubrum terrestre
  • **** Halorubrum tibetense
  • **** Halorubrum trapanicum
  • **** Halorubrum vacuolatum
  • **** Halorubrum yunnanense
  • **** Halorubrum xinjiangense
  • *** Halorussus
  • **** Halorussus amylolyticus
  • **** Halorussus rarus
  • **** Halorussus ruber
  • *** "Halosiccatus"
  • **** "Halosiccatus urmianus"
  • *** Halosimplex
  • **** Halosimplex carlsbadense
  • **** Halosimplex litoreum
  • **** Halosimplex pelagicum
  • **** Halosimplex rubrum
  • *** Halostagnicola
  • **** Halostagnicola alkaliphila
  • **** Halostagnicola bangensis
  • **** Halostagnicola kamekurae
  • **** Halostagnicola larsenii
  • *** "Halostella"
  • **** "Halostella salina"
  • *** Haloterrigena
  • **** Haloterrigena daqingensis
  • **** Haloterrigena hispanica
  • **** Haloterrigena jeotgali
  • **** Haloterrigena limicola
  • **** Haloterrigena longa
  • **** Haloterrigena saccharevitans
  • **** Haloterrigena salina
  • **** Haloterrigena thermotolerans
  • **** Haloterrigena turkmenica
  • *** Halovarius
  • **** Halovarius luteus
  • *** Halovenus
  • **** Halovenus aranensis
  • **** Halovenus rubra
  • **** Halovenus salina
  • *** Halovivax
  • **** Halovivax asiaticus
  • **** Halovivax cerinus
  • **** Halovivax limisalsi
  • **** Halovivax ruber
  • *** Natrialba
  • **** Natrialba aegyptia
  • **** Natrialba asiatica
  • **** Natrialba chahannaoensis
  • **** Natrialba hulunbeirensis
  • **** Natrialba magadii
  • **** Natrialba taiwanensis
  • *** Natribaculum
  • **** Natribaculum breve
  • **** Natribaculum longum
  • *** Natrinema
  • **** Natrinema altunense
  • **** Natrinema ejinorense
  • **** Natrinema gari
  • **** Natrinema pallidum
  • **** Natrinema pellirubrum
  • **** Natrinema salaciae
  • **** Natrinema versiforme
  • *** Natronoarchaeum
  • **** Natronoarchaeum mannanilyticum
  • **** Natronoarchaeum philippinense
  • **** Natronoarchaeum rubrum
  • *** Natronobacterium
  • **** Natronobacterium gregoryi
  • **** Natronobacterium texcoconense
  • *** Natronococcus
  • **** Natronococcus amylolyticus
  • **** Natronococcus jeotgali
  • **** Natronococcus occultus
  • **** Natronococcus roseus
  • *** Natronolimnobius
  • **** Natronolimnobius baerhuensis
  • **** Natronolimnobius innermongolicus
  • *** Natronomonas
  • **** Natronomonas gomsonensis
  • **** Natronomonas moolapensis
  • **** Natronomonas pharaonis
  • *** Natronorubrum
  • **** Natronorubrum aibiense
  • **** Natronorubrum bangense
  • **** Natronorubrum sediminis
  • **** Natronorubrum sulfidifaciens
  • **** Natronorubrum texcoconense
  • **** Natronorubrum tibetense
  • *** Salarchaeum
  • **** Salarchaeum japonicum
  • *** Salinarchaeum
  • **** Salinarchaeum laminariae
  • *** Salinigranum
  • **** Salinigranum rubrum
  • *** Salinirubrum
  • **** Salinirubrum litoreum
non-valid
  • "Halorubrum sfaxense"
  • "Candidatus Halobonum tyrrellensis
  • "Candidatus Haloectosymbiotes riaformosensis"
  • "Halopiger djelfamassiliensis sp"
  • "Halopiger goleamassiliensis"
  • "Halorubrum tropicale"
  • "Haloarcula rubripromontorii"
  • "Halobacterium hubeiense"
Classification of Gupta et al.
Halobacteriales
  Haladaptatus, Halalkalicoccus, Haloarchaeobius, Halarchaeum, Halobacterium, Halocalculus, Halorubellus, Halorussus, "Halosiccatus", Halovenus, Natronoarchaeum, Natronomonas, Salarchaeum.
  Halapricum, Haloarcula, Halomicroarcula, Halomicrobium, Halorientalis, Halorhabdus, Halosimplex.
  Halococcus.
Haloferacales
  Halabellus, Haloferax, Halogeometricum,, Halopelagius, Haloplanus, Haloquadratum, Halosarcina.
  Halobaculum,, Halohasta, Halolamina, Halonotius, Halopenitus, Halorubrum, Salinigranum.
Natrialbales
  Halobiforma, Halopiger, Halostagnicola, Haloterrigena, Halovarius, Halovivax, Natrialba, Natribaculum, Natronobacterium, Natronococcus, Natronolimnobius, Natronorubrum, Salinarchaeum.

Living environment

Haloarchaea require salt concentrations in excess of 2 M to grow, and optimal growth usually occurs at much higher concentrations, typically 20–25%. However, Haloarchaea can grow up to saturation.
Haloarchaea are found mainly in hypersaline lakes and solar salterns. Their high densities in the water often lead to pink or red colourations of the water. Some of them live in underground rock salt deposits, including one from middle-late Eocene. Some even older ones from more than 250 million years ago have been reported.

Adaptations to environment

Haloarchaea can grow at an aw close to 0.75, yet a water activity lower than 0.90 is inhibitory to most microbes. The number of solutes causes osmotic stress on microbes, which can cause cell lysis, unfolding of proteins and inactivation of enzymes when there is a large enough imbalance. Haloarchaea combat this by retaining compatible solutes such as potassium chloride in their intracellular space to allow them to balance osmotic pressure. Retaining these salts is referred to as the “salt-in” method where the cell accumulates a high internal concentration of potassium. Because of the elevated potassium levels, haloarchaea have specialized proteins that have a highly negative surface charge to tolerate high potassium concentrations.
Haloarchaea have adapted to use glycerol as a carbon and energy source in catabolic processes, which is often present in high salt environments due to Dunaliella species that produce glycerol in large quantities.

Phototrophy

is used to absorb light, which provides energy to transport protons across the cellular membrane. The concentration gradient generated from this process can then be used to synthesize ATP. Many haloarchaea also possess related pigments, including halorhodopsin, which pumps chloride ions in the cell in response to photons, creating a voltage gradient and assisting in the production of energy from light. The process is unrelated to other forms of photosynthesis involving electron transport, however, and haloarchaea are incapable of fixing carbon from carbon dioxide. Early evolution of retinal proteins has been proposed as the purple Earth hypothesis.

Cellular shapes

Haloarchaea are often considered pleomorphic, or able to take on a range of shapes—even within a single species. This makes identification by microscopic means difficult, and it is now more common to use gene sequencing techniques for identification instead.
One of the more unusually shaped Haloarchaea is the "Square Haloarchaeon of Walsby". It was classified in 2004 using a very low nutrition solution to allow growth along with a high salt concentration, square in shape and extremely thin. This shape is probably only permitted by the high osmolarity of the water, permitting cell shapes that would be difficult, if not impossible, under other conditions.

As exophiles

Haloarchaea have been proposed as a kind of life that could live on Mars; since the Martian atmosphere has a pressure below the triple point of water, freshwater species would have no habitat on the Martian surface. The presence of high salt concentrations in water lowers its freezing point, in theory allowing for halophiles to exist in saltwater on Mars.

Journals