Bacterial microcompartment
Encyclopedia
Bacterial microcompartments are widespread bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

l organelle
Organelle
In cell biology, an organelle is a specialized subunit within a cell that has a specific function, and is usually separately enclosed within its own lipid bilayer....

s that are made of a protein shell that surrounds and encloses various enzymes. These compartments are typically about 100-200 nanometre
Nanometre
A nanometre is a unit of length in the metric system, equal to one billionth of a metre. The name combines the SI prefix nano- with the parent unit name metre .The nanometre is often used to express dimensions on the atomic scale: the diameter...

s across and made of interlocking proteins. They do not contain lipid
Lipid
Lipids constitute a broad group of naturally occurring molecules that include fats, waxes, sterols, fat-soluble vitamins , monoglycerides, diglycerides, triglycerides, phospholipids, and others...

s since they are not surrounded by a membrane
Cell membrane
The cell membrane or plasma membrane is a biological membrane that separates the interior of all cells from the outside environment. The cell membrane is selectively permeable to ions and organic molecules and controls the movement of substances in and out of cells. It basically protects the cell...

. Protein-enclosed compartments are also found in eukaryotes, such as the mysterious vault complex
Vault (organelle)
The vault or vault cytoplasmic ribonucleoprotein is a eukaryotic organelle whose function is not fully understood. Discovered and successfully isolated by cell biologist Nancy Kedersha and biochemist Leonard Rome of the UCLA School of Medicine in the 1980s, vaults are cytoplasmic organelles which...

.

Protein families forming the microcompartment shell

The shells of diverse microcompartments are composed of members of three protein families: the BMC domain protein family, the inconsistently named CsoS4 / CcmL / EutN / OrfAB family, and the encapsulins/linocin-like proteins.

The BMC protein family

In microcompartment shells that have been studied, the major constituents are proteins belonging to the Bacterial Micro-Compartment (BMC) family. The crystal structures of a number of BMC proteins have been determined and invariably reveal assembly into cyclical hexamers with a small pore in the center.

The CsoS4 family

Recent structures have revealed either a pentameric or hexameric structure in this family. In icosahedral or quasi-icosahedral carboxysomes, it is likely that the pentameric form is positioned at the vertices.

Encapsulins

Encapsulins are a large and widely-distributed family of proteins and are present in most bacteria and have been identified in Candidatus methanoregula
Candidatus
Candidatus is in scientific classification a component of the taxonomic name for a bacterium that cannot be maintained in a Bacteriology Culture Collection. It is an interim taxonomic status for noncultivable organisms. An example would be "Candidatus Phytoplasma allocasuarinae"...

, a species of archaea
Archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...

. They were originally called linocin-like proteins and thought to be a group of bacterial antibiotics, since they showed bacteriostatic activity in culture. However, structural analysis showed these to form a spherical nanocompartment that contains enzymes involved in the defenses against oxidative stress
Oxidative stress
Oxidative stress represents an imbalance between the production and manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage...

.

Types

A recent survey indicated seven different metabolic systems encapsulated by microcompartment shells. Three are characterized:

Carboxysomes

Carboxysome
Carboxysome
Carboxysomes are bacterial microcompartments that contain enzymes involved in carbon fixation. Carboxysomes are made of polyhedral protein shells about 80 to 140 nanometres in diameter. These compartments are thought to concentrate carbon dioxide to overcome the inefficiency of RuBisCo - the...

s encapsulate RuBisCo
RuBisCO
Ribulose-1,5-bisphosphate carboxylase oxygenase, commonly known by the shorter name RuBisCO, is an enzyme involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is converted by plants to energy-rich molecules such as glucose. RuBisCo is an abbreviation...

 and carbonic anhydrase
Carbonic anhydrase
The carbonic anhydrases form a family of enzymes that catalyze the rapid interconversion of carbon dioxide and water to bicarbonate and protons , a reversible reaction that occurs rather slowly in the absence of a catalyst...

 in carbon-fixing bacteria as part of a carbon concentrating mechanism.

Pdu microcompartments

Some bacteria can used 1,2-propanediol as a carbon source. They express a microcompartment to encapsulate a number of enzymes used in this pathway. The Pdu compartment is constructed by a set of 21 genes in a single chromosomal locus. These genes are sufficient for assembly of the microcompartment since they can be transferred between bacteria and will produce a functional structure in the recipient.

Eut microcompartments

EUT microcompartments are proposed to form in Salmonella
Salmonella
Salmonella is a genus of rod-shaped, Gram-negative, non-spore-forming, predominantly motile enterobacteria with diameters around 0.7 to 1.5 µm, lengths from 2 to 5 µm, and flagella which grade in all directions . They are chemoorganotrophs, obtaining their energy from oxidation and reduction...

and other Enterobacteriaceae
Enterobacteriaceae
The Enterobacteriaceae is a large family of bacteria that includes many of the more familiar pathogens, such as Salmonella, Escherichia coli, Yersinia pestis, Klebsiella and Shigella. This family is the only representative in the order Enterobacteriales of the class Gammaproteobacteria in the...

 species, and are involved in the metabolism of ethanolamine
Ethanolamine
Ethanolamine, also called 2-aminoethanol or monoethanolamine , is an organic chemical compound that is both a primary amine and a primary alcohol . Like other amines, monoethanolamine acts as a weak base...

.

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