Osmoprotectants or compatible solutes are small organic molecules with neutral charge and low toxicity at high concentrations that act as
osmolyte
Osmolytes are low-molecular weight organic compounds that influence the properties of biological fluids. Their primary role is to maintain the integrity of cells by affecting the viscosity, melting point, and ionic strength of the aqueous solution. ...
s and help organisms survive extreme
osmotic stress
Osmotic shock or osmotic stress is physiologic dysfunction caused by a sudden change in the solute concentration around a cell, which causes a rapid change in the movement of water across its cell membrane. Under hypertonic conditions - conditio ...
.
Osmoprotectants can be placed in three chemical classes: betaines and associated molecules, sugars and polyols, and amino acids. These molecules accumulate in cells and balance the osmotic difference between the cell's surroundings and the
cytosol
The cytosol, also known as cytoplasmic matrix or groundplasm, is one of the liquids found inside cells (intracellular fluid (ICF)). It is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondri ...
.
In plants, their accumulation can increase survival during stresses such as drought. In extreme cases, such as in
bdelloid rotifers
Bdelloidea (Greek ''βδέλλα'', ''bdella'', "leech") is a class of rotifers found in freshwater habitats all over the world. There are over 450 described species of bdelloid rotifers (or 'bdelloids'), distinguished from each other mainl ...
,
tardigrades
Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbär ...
,
brine shrimp
''Artemia'' is a genus of aquatic crustaceans also known as brine shrimp. It is the only genus in the family Artemiidae. The first historical record of the existence of ''Artemia'' dates back to the first half of the 10th century AD from Urmia L ...
, and
nematodes
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
, these molecules can allow cells to survive being completely dried out and let them enter a state of suspended animation called
cryptobiosis.
Intracellular osmoprotectant concentrations are regulated in response to environmental conditions such as
osmolarity
Osmotic concentration, formerly known as osmolarity, is the measure of solute concentration, defined as the number of osmoles (Osm) of solute per litre (L) of solution (osmol/L or Osm/L). The osmolarity of a solution is usually expressed as Osm/L ...
and temperature via regulation of specific
transcription factors
In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The func ...
and
transporters. They have been shown to play a protective role by maintaining enzyme activity through freeze-thaw cycles and at higher temperatures. It is currently believed that they function by stabilizing protein structures by promoting preferential exclusion from the water layers on the surface of hydrated proteins. This favors the native conformation and displaces inorganic salts that would otherwise cause misfolding.
Role
Compatible solutes have a functional role in agriculture. In high stress conditions, such as drought or high salinity, plants that naturally create or take up osmoprotectants show increased survival rates. By inducing expression or uptake of these molecules in crops in which they are naturally not present, there is an increase in the areas in which they are able to be grown. One documented reason for increased growth is regulation of toxic
reactive oxygen species
In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen.
The reduction of molecular oxygen () p ...
(ROS). In high salinity ROS production is stimulated by the
photosystems
Photosystems are functional and structural units of protein complexes involved in photosynthesis. Together they carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. Photosystems ...
of the plant. Osmoprotectants can prevent the photosystem-salt interactions, reducing ROS production. For these reasons, introduction of biosynthetic pathways which result in the creation of osmoprotectants in crops is a current area of research, but inducing expression at significant amounts is currently posing a barrier in this area of research.
Osmoprotectants are also important for the maintenance of top soil bacteria populations. Desiccation of top soils results in increased salinity. In these situations, the soil microbes increase the concentration of these molecule in their cytoplasm into the molar range allowing them to persist until conditions approve.
In extreme cases, osmoprotectants allow cells to enter cryptobiosis. In this state the cytosol and osmoprotectants become a glass-like solid that helps stabilize
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s and
cell membrane
The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
s from the damaging effects of desiccation.
Additionally, osmoprotectants provide a method to regulate gene expression in response to environmental osmolarity. The presence of compatible solutes even in small concentrations has been shown to affect gene expression. Their effects range from inducing production of more compatible solutes to regulating components involved in infection, such as
phospholipase
A phospholipase is an enzyme that hydrolyzes phospholipids into fatty acids and other lipophilic substances. Acids trigger the release of bound calcium from cellular stores and the consequent increase in free cytosolic Ca2+, an essential step in ...
C in ''
Pseudomonas aeruginosa
''Pseudomonas aeruginosa'' is a common encapsulated, gram-negative, aerobic–facultatively anaerobic, rod-shaped bacterium that can cause disease in plants and animals, including humans. A species of considerable medical importance, ''P. aerugi ...
''.
See also
*
Ectoine
Ectoine (1,4,5,6-tetrahydro-2-methyl-4-pyrimidinecarboxylic acid) is a natural compound found in several species of bacteria. It is a compatible solute which serves as a protective substance by acting as an osmolyte and thus helps organisms surv ...
*
Osmoregulation
Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
*
Proline
References
Cell biology
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