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Halotolerance is the
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
of living
organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...
s to conditions of high
salinity Salinity () is the saltiness or amount of salt (chemistry), salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensio ...
. Halotolerant species tend to live in areas such as hypersaline lakes,
coastal dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
, saline
desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...
s,
salt marsh A salt marsh, saltmarsh or salting, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastal intertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. I ...
es, and inland salt
sea A sea is a large body of salt water. There are particular seas and the sea. The sea commonly refers to the ocean, the interconnected body of seawaters that spans most of Earth. Particular seas are either marginal seas, second-order section ...
s and springs.
Halophile A halophile (from the Greek word for 'salt-loving') is an extremophile that thrives in high salt In common usage, salt is a mineral composed primarily of sodium chloride (NaCl). When used in food, especially in granulated form, it is more ...
s are also organisms that not only live in highly saline environments but also ''require'' the
salinity Salinity () is the saltiness or amount of salt (chemistry), salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensio ...
to survive. Halotolerant organisms on the other hand (belonging to different domains of life) can grow under saline conditions, but do not require elevated concentrations of salt for growth.
Halophyte A halophyte is a salt-tolerant plant that grows in soil or waters of high salinity, coming into contact with saline water through its roots or by salt spray, such as in saline semi-deserts, mangrove swamps, marshes and sloughs, and seashores. ...
s are salt-tolerant higher plants. Halotolerant microorganisms are of considerable biotechnological interest.


Applications

Fields of scientific research relevant to halotolerance include
biochemistry Biochemistry, or biological chemistry, is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology, a ...
,
molecular biology Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
,
cell biology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living an ...
,
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
,
ecology Ecology () is the natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...
, and
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
. An understanding of halotolerance can be applicable to areas such as arid-zone agriculture,
xeriscaping Xeriscaping is the process of Garden design, landscaping, or gardening, that reduces or eliminates the need for irrigation. It is promoted in regions that do not have accessible, plentiful, or reliable supplies of fresh water and has gained acce ...
,
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
(of fish or algae), bioproduction of desirable compounds (such as
phycobiliprotein Phycobiliproteins are water-soluble proteins present in cyanobacteria and certain algae (rhodophytes, cryptomonads, glaucocystophytes). They capture light energy, which is then passed on to chlorophylls during photosynthesis. Phycobiliproteins are ...
s or carotenoids) using seawater to support growth, or remediation of salt-affected soils. In addition, many environmental stressors involve or induce osmotic changes, so knowledge gained about halotolerance can also be relevant to understanding tolerance to extremes in moisture or temperature. Goals of studying halotolerance include increasing the agricultural productivity of lands affected by soil salination or where only saline water is available. Conventional agricultural species could be made more halotolerant by gene transfer from naturally halotolerant species (by conventional breeding or
genetic engineering Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of Genetic engineering techniques, technologies used to change the genet ...
) or by applying treatments developed from an understanding of the mechanisms of halotolerance. In addition, naturally halotolerant plants or microorganisms could be developed into useful
agricultural Agriculture encompasses crop and livestock production, aquaculture, and forestry for food and non-food products. Agriculture was a key factor in the rise of sedentary human civilization, whereby farming of domesticated species created f ...
crops or
fermentation Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic molecules, such as glucose or other sugars, are catabolized and reduce ...
organisms.


Cellular functions in halophytes

Tolerance of high salt conditions can be obtained through several routes. High levels of salt entering the plant can trigger ionic imbalances which cause complications in respiration and photosynthesis, leading to reduced rates of growth, injury and death in severe cases. To be considered tolerant of saline conditions, the
protoplast Protoplast (), is a biology, biological term coined by Johannes von Hanstein, Hanstein in 1880 to refer to the entire cell, excluding the cell wall. Protoplasts can be generated by stripping the cell wall from plant, bacterium, bacterial, or f ...
must show methods of balancing the toxic and osmotic effects of the increased salt concentrations. Halophytic vascular plants can survive on soils with salt concentrations around 6%, or up to 20% in extreme cases ( ocean salinity is around 3.5%). Tolerance of such conditions is reached through the use of stress proteins and compatible cytoplasm osmotic solutes. To exist in such conditions, halophytes tend to be subject to the uptake of high levels of salt into their cells, and this is often required to maintain an osmotic potential lower than that of the soil to ensure water uptake. High salt concentrations within the cell can be damaging to sensitive organelles such as the chloroplast, so sequestration of salt is seen. Under this action, salt is stored within the
vacuole A vacuole () is a membrane-bound organelle which is present in Plant cell, plant and Fungus, fungal Cell (biology), cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water ...
to protect such delicate areas. If high salt concentrations are seen within the vacuole, a high concentration gradient will be established between the vacuole and the cytoplasm, leading to high levels of energy investment to maintain this state. Therefore, the accumulation of compatible cytoplasmic osmotic solutes can be seen to prevent this situation from occurring.
Amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...
s such as proline accumulate in halophytic Brassica species, quaternary ammonium bases such as Glycine Betaine and sugars have been shown to act in this role within halophytic members of
Chenopodiaceae Amaranthaceae ( ) is a family of flowering plants commonly known as the amaranth family, in reference to its type (biology), type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 ...
and members of
Asteraceae Asteraceae () is a large family (biology), family of flowering plants that consists of over 32,000 known species in over 1,900 genera within the Order (biology), order Asterales. The number of species in Asteraceae is rivaled only by the Orchi ...
show the buildup of cyclites and soluble sugars. The buildup of these compounds allow for the balancing of the osmotic effect while preventing the establishment of toxic concentrations of salt or requiring the maintenance of high concentration gradients.


Bacterial halotolerance

The extent of halotolerance varies widely amongst different species of bacteria. A number of
cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
are halotolerant; an example location of occurrence for such cyanobacteria is in the Makgadikgadi Pans, a large hypersaline lake in
Botswana Botswana, officially the Republic of Botswana, is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory part of the Kalahari Desert. It is bordered by South Africa to the sou ...
.


Fungal halotolerance

Fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
from habitats with high concentration of salt are mostly halotolerant (i.e. they do not require salt for growth) and not halophilic. Halophilic fungi are a rare exception. Halotolerant fungi constitute a relatively large and constant part of hypersaline environment communities, such as those in the solar salterns. Well studied examples include the yeast '' Debaryomyces hansenii ''and black yeasts '' Aureobasidium pullulans'' and '' Hortaea werneckii''. The latter can grow in media without salt, as well as in almost saturated NaCl solutions. To emphasize this unusually wide
adaptability Adaptability ( "fit to, adjust") is a feature of a system or of a process. This word has been put to use as a specialised term in different disciplines and in business operations. Word definitions of adaptability as a specialised term differ littl ...
, some authors describe ''H. werneckii ''as "extremely halotolerant".


See also

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References

{{Animalosmo Environmental microbiology Geomicrobiology Microbial growth and nutrition de:Halophil