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''Caudovirales'' is an order of
virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsk ...
es known as the tailed
bacteriophage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bac ...
s (''cauda'' is Latin for "tail"). Under the Baltimore classification scheme, the ''Caudovirales'' are group I viruses as they have double stranded DNA (dsDNA) genomes, which can be anywhere from 18,000 base pairs to 500,000 base pairs in length. The virus particles have a distinct shape; each virion has an icosahedral head that contains the viral genome, and is attached to a flexible tail by a connector protein. The order encompasses a wide range of viruses, many containing genes of similar nucleotide sequence and function. However, some tailed bacteriophage genomes can vary quite significantly in nucleotide sequence, even among the same genus. Due to their characteristic structure and possession of potentially homologous genes, it is believed these bacteriophages possess a common origin. There are 14 families, 73 subfamilies, 927 genera, and 2,814 species in the order. This makes ''Caudovirales'' the most populous order among viruses, accounting for approximately 30% of all recognized virus species and nearly half of all virus genera.


Infection

Upon encountering a host bacterium, the tail section of the virion binds to receptors on the cell surface and delivers the DNA into the cell by use of an injectisome-like mechanism (an injectisome is a nanomachine that evolved for the delivery of proteins by type III secretion). The tail section of the virus punches a hole through the bacterial cell wall and plasma membrane and the genome passes down the tail into the cell. Once inside the genes are expressed from transcripts made by the host machinery, using host
ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to fo ...
s. Typically, the genome is replicated by use of
concatemer A concatemer is a long continuous DNA molecule that contains multiple copies of the same DNA sequence linked in series. These polymeric molecules are usually copies of an entire genome linked end to end and separated by ''cos'' sites (a protein b ...
s, in which overlapping segments of DNA are made, and then put together to form the whole genome.


Assembly and maturation

Viral
capsid A capsid is the protein shell of a virus, enclosing its genetic material. It consists of several oligomeric (repeating) structural subunits made of protein called protomers. The observable 3-dimensional morphological subunits, which may or ma ...
proteins come together to form a precursor prohead, into which the genome enters. Once this has occurred, the prohead undergoes maturation by cleavage of capsid subunits to form an icosahedral
phage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bacter ...
head with 5-fold symmetry. After the head maturation, the tail is joined in one of two ways: Either the tail is constructed separately, and joined with the connector, or the tail is constructed directly onto the phage head. The tails consist of helix based proteins with 6-fold symmetry. After maturation of virus particles, the cell is lysed by
lysin Lysins, also known as endolysins or murein hydrolases, are hydrolytic enzymes produced by bacteriophages in order to cleave the host's cell wall during the final stage of the lytic cycle. Lysins are highly evolved enzymes that are able to target ...
s, holins, or a combination of the two.


Taxonomy

Because of the lack of homology between the amino acid and DNA sequences of these viruses these factors are precluded from being used as taxonomic markers as is common for other organisms, the three families discussed below (''
Myoviridae ''Myoviridae'' is a family of bacteriophages in the order '' Caudovirales''. Bacteria and archaea serve as natural hosts. There are 625 species in this family, assigned to eight subfamilies and 217 genera. Subdivisions The subfamily ''Tevenvirina ...
'', '' Podoviridae'', and '' Siphoviridae'') are defined on the basis of morphology. This classification scheme was originated by Bradley in 1969 and has been extended since. All viruses in this order have icosahedral or oblate heads but differ in the length and contractile abilities of their tails. The ''Myoviridae'' have long tails that are contractile; the ''Podoviridae'' have short noncontractile tails; and the ''Siphoviridae'' have long noncontractile tails. ''Siphoviridae'' constitute the majority of the known tailed viruses. Bradley referred to what is now known as the ''Myoviridae'' as type A, ''Siphoviridae'' as type B, and the ''Podoviridae'' as type C. He also divided his groups on the basis of head morphology: Within group A, A1 have small isometric heads; A2 have prolate heads; and A3 have elongated heads. Within groups B and C, numbers were similarly assigned: B1 and C1 have small isometric heads; B2 and C2 have prolate heads; and B3 and C3 have elongated heads. In addition the aforementioned three families, a total of 14 families and one ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' genus are assigned to the order: * '' Ackermannviridae'' * '' Autographiviridae'' * '' Chaseviridae'' * '' Demerecviridae'' * '' Drexlerviridae'' * '' Guelinviridae'' * '' Herelleviridae'' * ''
Myoviridae ''Myoviridae'' is a family of bacteriophages in the order '' Caudovirales''. Bacteria and archaea serve as natural hosts. There are 625 species in this family, assigned to eight subfamilies and 217 genera. Subdivisions The subfamily ''Tevenvirina ...
'' * '' Podoviridae'' * '' Rountreeviridae'' * '' Salasmaviridae'' * '' Schitoviridae'' * '' Siphoviridae'' * '' Zobellviridae'' * '' Lilyvirus'', the unassigned genus


Bacteriophage evolution

Bacteriophages occur in over 195 bacterial or archaeal genera. They arose repeatedly in different hosts and there are at least 11 separate lines of descent. Over 6300 bacteriophages have been examined in the electron microscope since 1959. Of these, more than 96 percent have tails. Of the tailed phages, about 57 percent have long, noncontractile tails (''Siphoviridae''). Tailed phages appear to be monophyletic and are the oldest known virus group.


See also

*
WO virus WO virus is bacteriophage virus that infects bacteria of the genus ''Wolbachia'', which it is named after. This virus is notable for carrying DNA related to the black widow spider toxin gene, becoming an example of a bacteriophage with animal-li ...


References


Further reading

* * {{Taxonbar, from=Q6521 Archaeal viruses Bacteriophages Virus orders