Pilus
Encyclopedia
A pilus is a hairlike appendage found on the surface of many bacteria. The terms pilus and fimbria
(Latin for 'thread' or 'fiber'; plural: fimbriae) can be used interchangeably, although some researchers reserve the term pilus for the appendage required for bacterial conjugation
. All pili are primarily composed of oligomer
ic pilin
proteins.
Dozens of these structures can exist on the bacteria. Some bacterial virus
es or bacteriophage
s attach to receptors
on pili at the start of their reproductive cycle.
Pili are antigenic. They are also fragile and constantly replaced, sometimes with pili of different composition, resulting in altered antigenicity. Specific host responses to old pili structure are not effective on the new structure. Recombination genes of pili code for variable (V) and constant (C) regions of the pili (similar to immunoglobulin diversity).
between bacteria, in the process of bacterial conjugation
. They are sometimes called "sex pili", in analogy to sexual reproduction
, because they allow for the exchange of genes via the formation of "mating pairs". Perhaps the most well-studied is the F pilus of Escherichia coli
, encoded by the F plasmid or fertility factor.
A pilus is typically 6 to 7 nm in diameter. During conjugation, a pilus emerging from donor bacterium ensnares the recipient bacterium, draws it in close, and eventually triggers the formation of a mating bridge, which establishes direct contact and the formation of a controlled pore that allows transfer of DNA from the donor to the recipient. Typically, the DNA transferred consists of the genes required to make and transfer pili (often encoded on a plasmid
), and so is a kind of selfish DNA
; however, other pieces of DNA are often co-transferred and this can result in dissemination of genetic traits, such as antibiotic resistance
, among a bacterial population. Not all bacteria can make conjugative pili, but conjugation can occur between bacteria of different species.
forces. The external ends of the pili adhere to a solid substrate, either the surface to which the bacteria are attached or to other bacteria, and when the pilus contracts, it pulls the bacteria forward, like a grappling hook. Movement produced by type IV pili is typically jerky, and so it is called twitching motility, as distinct from other forms of bacterial motility, such as motility produced by flagella. However, some bacteria, for example Myxococcus xanthus
, exhibit gliding motility. Bacterial type IV pilins are similar in structure to the component flagellins of Archaeal flagella.
. A fimbria
is a short pilus that is used to attach the bacterium to a surface. Fimbriae are either located at the poles of a cell, or are evenly spread over its entire surface. Mutant
bacteria that lack fimbriae cannot adhere to their usual target surfaces and, thus, cannot cause disease
s.
Some fimbriae can contain lectins. The lectins are necessary to adhere to target cells because they can recognize oligosaccharide
units on the surface of these target cells. Other fimbriae bind to components of the extracellular matrix
.
Fimbriae are found in both Gram-negative
and Gram-positive
bacteria. In Gram-positive bacteria, the pilin subunits are covalently linked.
Fimbria (bacteriology)
In bacteriology, a Cilli is a proteinaceous appendage in many Gram-negative and some Gram-positive bacteria that is thinner and shorter than a flagellum. This appendage ranges from 3-10 nanometers in diameter and can be up to several micrometers long...
(Latin for 'thread' or 'fiber'; plural: fimbriae) can be used interchangeably, although some researchers reserve the term pilus for the appendage required for bacterial conjugation
Bacterial conjugation
Bacterial conjugation is the transfer of genetic material between bacterial cells by direct cell-to-cell contact or by a bridge-like connection between two cells...
. All pili are primarily composed of oligomer
Oligomer
In chemistry, an oligomer is a molecule that consists of a few monomer units , in contrast to a polymer that, at least in principle, consists of an unlimited number of monomers. Dimers, trimers, and tetramers are oligomers. Many oils are oligomeric, such as liquid paraffin...
ic pilin
Pilin
Pilin refers to a class of fibrous proteins that are found in pilus structures in bacteria. Bacterial pili are used in the exchange of genetic material during bacterial conjugation, and a short pilus called a fimbrium is used as a cell adhesion mechanism. Although not all bacteria have pili or...
proteins.
Dozens of these structures can exist on the bacteria. Some bacterial virus
Virus
A virus is a small infectious agent that can replicate only inside the living cells of organisms. Viruses infect all types of organisms, from animals and plants to bacteria and archaea...
es or bacteriophage
Bacteriophage
A bacteriophage is any one of a number of viruses that infect bacteria. They do this by injecting genetic material, which they carry enclosed in an outer protein capsid...
s attach to receptors
Receptor (biochemistry)
In biochemistry, a receptor is a molecule found on the surface of a cell, which receives specific chemical signals from neighbouring cells or the wider environment within an organism...
on pili at the start of their reproductive cycle.
Pili are antigenic. They are also fragile and constantly replaced, sometimes with pili of different composition, resulting in altered antigenicity. Specific host responses to old pili structure are not effective on the new structure. Recombination genes of pili code for variable (V) and constant (C) regions of the pili (similar to immunoglobulin diversity).
Conjugative pili
Conjugative pili allow the transfer of DNADNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
between bacteria, in the process of bacterial conjugation
Bacterial conjugation
Bacterial conjugation is the transfer of genetic material between bacterial cells by direct cell-to-cell contact or by a bridge-like connection between two cells...
. They are sometimes called "sex pili", in analogy to sexual reproduction
Sexual reproduction
Sexual reproduction is the creation of a new organism by combining the genetic material of two organisms. There are two main processes during sexual reproduction; they are: meiosis, involving the halving of the number of chromosomes; and fertilization, involving the fusion of two gametes and the...
, because they allow for the exchange of genes via the formation of "mating pairs". Perhaps the most well-studied is the F pilus of Escherichia coli
Escherichia coli
Escherichia coli is a Gram-negative, rod-shaped bacterium that is commonly found in the lower intestine of warm-blooded organisms . Most E. coli strains are harmless, but some serotypes can cause serious food poisoning in humans, and are occasionally responsible for product recalls...
, encoded by the F plasmid or fertility factor.
A pilus is typically 6 to 7 nm in diameter. During conjugation, a pilus emerging from donor bacterium ensnares the recipient bacterium, draws it in close, and eventually triggers the formation of a mating bridge, which establishes direct contact and the formation of a controlled pore that allows transfer of DNA from the donor to the recipient. Typically, the DNA transferred consists of the genes required to make and transfer pili (often encoded on a plasmid
Plasmid
In microbiology and genetics, a plasmid is a DNA molecule that is separate from, and can replicate independently of, the chromosomal DNA. They are double-stranded and, in many cases, circular...
), and so is a kind of selfish DNA
Selfish DNA
Selfish DNA refers to those sequences of DNA which, in their purest form, have two distinct properties: the DNA sequence spreads by forming additional copies of itself within the genome; and it makes no specific contribution to the reproductive success of its host organism.This idea was sketched...
; however, other pieces of DNA are often co-transferred and this can result in dissemination of genetic traits, such as antibiotic resistance
Antibiotic resistance
Antibiotic resistance is a type of drug resistance where a microorganism is able to survive exposure to an antibiotic. While a spontaneous or induced genetic mutation in bacteria may confer resistance to antimicrobial drugs, genes that confer resistance can be transferred between bacteria in a...
, among a bacterial population. Not all bacteria can make conjugative pili, but conjugation can occur between bacteria of different species.
Type IV pili
Some pili, called type IV pili, generate motileMotility
Motility is a biological term which refers to the ability to move spontaneously and actively, consuming energy in the process. Most animals are motile but the term applies to single-celled and simple multicellular organisms, as well as to some mechanisms of fluid flow in multicellular organs, in...
forces. The external ends of the pili adhere to a solid substrate, either the surface to which the bacteria are attached or to other bacteria, and when the pilus contracts, it pulls the bacteria forward, like a grappling hook. Movement produced by type IV pili is typically jerky, and so it is called twitching motility, as distinct from other forms of bacterial motility, such as motility produced by flagella. However, some bacteria, for example Myxococcus xanthus
Myxococcus xanthus
Myxococcus xanthus colonies exist as a self-organized, predatory, saprotrophic, single-species biofilm called a swarm. Myxococcus xanthus, which can be found almost ubiquitously in soil, are thin rod shaped, gram-negative cells that exhibit self-organizing behavior as a response to environmental...
, exhibit gliding motility. Bacterial type IV pilins are similar in structure to the component flagellins of Archaeal flagella.
Fimbriae
Attachment of bacteria to host surfaces is required for colonization during infection or to initiate formation of a biofilmBiofilm
A biofilm is an aggregate of microorganisms in which cells adhere to each other on a surface. These adherent cells are frequently embedded within a self-produced matrix of extracellular polymeric substance...
. A fimbria
Fimbria (bacteriology)
In bacteriology, a Cilli is a proteinaceous appendage in many Gram-negative and some Gram-positive bacteria that is thinner and shorter than a flagellum. This appendage ranges from 3-10 nanometers in diameter and can be up to several micrometers long...
is a short pilus that is used to attach the bacterium to a surface. Fimbriae are either located at the poles of a cell, or are evenly spread over its entire surface. Mutant
Mutant
In biology and especially genetics, a mutant is an individual, organism, or new genetic character, arising or resulting from an instance of mutation, which is a base-pair sequence change within the DNA of a gene or chromosome of an organism resulting in the creation of a new character or trait not...
bacteria that lack fimbriae cannot adhere to their usual target surfaces and, thus, cannot cause disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...
s.
Some fimbriae can contain lectins. The lectins are necessary to adhere to target cells because they can recognize oligosaccharide
Oligosaccharide
An oligosaccharide is a saccharide polymer containing a small number of component sugars, also known as simple sugars...
units on the surface of these target cells. Other fimbriae bind to components of the extracellular matrix
Extracellular matrix
In biology, the extracellular matrix is the extracellular part of animal tissue that usually provides structural support to the animal cells in addition to performing various other important functions. The extracellular matrix is the defining feature of connective tissue in animals.Extracellular...
.
Fimbriae are found in both Gram-negative
Gram-negative
Gram-negative bacteria are bacteria that do not retain crystal violet dye in the Gram staining protocol. In a Gram stain test, a counterstain is added after the crystal violet, coloring all Gram-negative bacteria with a red or pink color...
and Gram-positive
Gram-positive
Gram-positive bacteria are those that are stained dark blue or violet by Gram staining. This is in contrast to Gram-negative bacteria, which cannot retain the crystal violet stain, instead taking up the counterstain and appearing red or pink...
bacteria. In Gram-positive bacteria, the pilin subunits are covalently linked.