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bacteriophage

 
American Heritage Dictionary:

bac·te·ri·o·phage

(băk-tîr'ē-ə-fāj') pronunciation
n.
A virus that infects and lyses certain bacteria.

bacteriophagic bac·te'ri·o·phag'ic (-făj'ĭk) adj.
bacteriophagy bac·te'ri·oph'a·gy (-ŏf'ə-jē) n.

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Any of a group of usually complex viruses that infect bacteria. Discovered in the early 20th century, bacteriophages were used to treat human bacterial diseases such as bubonic plague and cholera but were not successful; they were abandoned with the advent of antibiotics in the 1940s. The rise of drug-resistant bacteria in the 1990s focused renewed attention on the therapeutic potential of bacteriophages. Thousands of varieties exist, each of which may infect only one or a few types of bacteria. The core of a bacteriophage's genetic material may be either DNA or RNA. On infecting a host cell, bacteriophages known as lytic or virulent phages release replicated viral particles by lysing (bursting) the host cell. Other types, known as lysogenic or temperate, integrate their nucleic acid into the host's chromosome to be replicated during cell division. During this time they are not virulent. The viral genome may later become active, initiating production of viral particles and destruction of the host cell. A.D. Hershey and Martha Chase used a bacteriophage in a famous 1952 experiment that supported the theory that DNA is the genetic material. Because bacteriophage genomes are small and because large quantities can be prepared in the laboratory, they are a favourite research tool of molecular biologists. Studies of phages have helped illuminate genetic recombination, nucleic acid replication, and protein synthesis.

For more information on bacteriophage, visit Britannica.com.

Any of the viruses that infect bacterial cells. They are discrete particles with dimensions from about 20 to about 200 nanometers. A given bacterial virus can infect only one or a few related species of bacteria; these constitute its host range. Bacteriophages consist of two essential components: nucleic acid, in which genetic information is encoded (this may be either ribonucleic acid or deoxyribonucleic acid), and a protein coat (capsid), which serves as a protective shell containing the nucleic acid and is involved in the efficiency of infection and the host range of the virus.

The description of a bacterial virus involves a study of its shape and dimensions by electron microscopy (see illustration), its host range, the serological properties of its capsid, the kind of nucleic acid it contains, and the characters of the plaques it forms on a given host. Both the nucleic acid and the capsid proteins are specific to the individual virus; in the case of the capsid proteins this specificity is the basis for serological identification of the virus.

Diagram of a T4 bacteriophage.
Diagram of a T4 bacteriophage.

The most striking form of phage infection is that in which all of the infected bacteria are destroyed in the process of the formation of new phage particles. This results in the clearing of a turbid liquid culture as the infected cells lyse. When lysis occurs in cells fixed as a lawn of bacteria growing on a solid medium, it produces holes, or areas of clearing, called plaques. These represent colonies of bacteriophage. The size and other properties of the plaque vary with individual viruses and host cells. See also Actinophage; Coliphage; Lysogeny; Lytic infection; Virus.


Viruses that attack bacteria, commonly known as phages. They pass through bacterial filters, and can be a cause of considerable trouble in bacterial cultures (for example milk starter cultures).

Columbia Encyclopedia:

bacteriophage

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bacteriophage (băktēr'ēəfāj'), virus that infects bacteria and sometimes destroys them by lysis, or dissolution of the cell. Bacteriophages, or phages, have a head composed of protein, an inner core of nucleic acid-either deoxyribonucleic acid (DNA) or ribonucleic acid (RNA)-and a hollow protein tail. A particular phage can usually infect only one or a few related species of bacteria; for example, coliphages are DNA-containing viruses that infect only the bacterium Escherichia coli.

A virus infects a bacterial cell by first attaching to the bacterial cell wall by its tail. In coliphages the tail is a complex protein structure consisting of a hollow contractile sheath, with a plate at the base that contains long protein fibers. The tail fibers fix the base plate to the specific receptor site on the bacterial cell wall, and the tail sheath contracts like a syringe, forcing the DNA that is inside the virus through the cell wall and cell membrane. The entire virus protein coat remains outside the bacterium.

The injected nucleic acid is the viral genetic material; it makes use of the bacterium's chemical energy and biosynthetic machinery to produce viral enzymes, as well as more phage nucleic acid. The viral proteins and nucleic acid molecules within the bacterial host assemble spontaneously into up to a hundred new phage particles. Eventually the bacterium lyses, releasing the particles. Lysis can be readily observed in bacteria growing on a solid medium, where groups of lysed cells appear as clear areas, or plaques.

Some DNA phages, called temperate phages, only lyse a small fraction of bacterial cells; in the remaining majority of the bacteria, the phage DNA becomes integrated into the bacterial chromosome and replicates along with it. In this state, known as lysogeny, the information contained in the viral nucleic acid is not expressed. A lysogenic bacterial culture can be treated with radiation or mutagens, inducing the cells to begin producing viruses and lyse. Lysogenic phages resemble bacterial genetic particles known as episomes. Incorporated phage genes are sometimes the source of the virulence of disease-causing bacteria.

The bacteriophage was discovered independently by the microbiologists F. W. Twort (1915) and Félix d'Hérelle (1917). The phages have been much used in the study of bacterial genetics and cellular control mechanisms largely because the bacterial hosts are so easily grown and infected with phage in the laboratory. Phages were also used in an attempt to destroy bacteria that cause epidemic diseases, but this approach was largely abandoned in the 1940s when antibacterial drugs became available. The possibility of "phage therapy" has recently attracted new interest among medical researchers, however, owing to the increasing threat posed by drug-resistant bacteria. In 2006 the Food and Drug Administration approved the use of bacteriophages that attack strains of Listeria as a food additive on ready-to-eat meat products.


Biology Q&A:

What is a bacteriophage?

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A bacteriophage, also called a phage, is a virus that infects bacteria. The term "bacteriophage" means "bacteria eater" (from the Greek word phagein, which means "to devour"). Phages consist of a long nucleic acid molecule (usually DNA) coiled within a polyhedral-shaped protein head. Many phages have a tail attached to the head. Fibers extending from the tail may be used to attach the virus to the bacterium.

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Wiley Dictionary of Flavors:

Bacteriophage

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A virus that attacks and consumes a bacterium. Dairy products are susceptible to these microorganisms. Usually phages are specific to certain organisms.

or phage or bacterial virus

any virus that can infect and multiply in a bacterium. Phages parasitize almost every group of prokaryotes. Commonly they consist of a core of nucleic acid enclosed within a protein coat, the capsid; additional components, e.g. lipid, occur in some phages. Depending on the type of phage, the nucleic acid may be either DNA (single-stranded or double-stranded) or RNA (single-stranded), and may be either linear or circular. Phages may be filamentous, polyhedral, or polyhedral and tailed; the tubular tails, to which one or more tubular tail fibres are attached in certain circumstances, are involved in attachment of the phages to the bacterial surface and for the injection of the DNA into the host cells. The smallest phages, e.g. φX174, are ≈25 nm in diameter, whereas tailed polyhedral phages, such as the T-even phages, are 200 — 250 nm long, while filamentous phages measure roughly 800 nm × 5 nm. In general, a given bacteriophage can infect only one particular species, or strain, or group of closely related strains, though a given strain may be susceptible to infection by a number of different phages. Bacteriophages can bring about transduction (def. 1) between bacterial cells. Compare cyanophage. See (bacterio)phage conversion, lysogeny, phage induction, phage typing, prophage, temperate phage, virulent phage.

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Or simply phage; a virus that infects bacteria often killing them by lysis; many varieties exist, and usually each attacks only one kind of bacteria. Some bacteriophages are widely used as cloning vectors and for determining DNA sequence. Virulent DNA bacteriophages in the T series adsorb to specific receptor sites on the bacterial cell wall and inject their DNA content into the bacterium. The viral DNA usurps the machinery of the cell for the replication of viral DNA and protein which is assembled into a crop of progeny phage which are released by lysis from the cell. Called also bacterial virus.

  • M13 b. — small rod-shaped, nonlytic, single-stranded DNA phage; used as a template in the Sangar dideoxy method for DNA sequence determination.
  • φX174 b. — prototype of a class of phage that are small, icosahedral single-stranded DNA viruses that code for only 10 to 12 proteins and are highly dependent on the host cell for their replication.
  • RNA b. — the genome is RNA instead of DNA; smallest known viruses, encode for only four proteins; they have contributed to basic studies of RNA.
  • temperate b's — typified by λ phage; have a similar lytic life cycle but in addition have an alternate life cycle whereby the injected DNA becomes integrated into the cell DNA where it remains stable, behaving as a cell gene. The integrated DNA is called prophage and the bacterial cell is said to be lysogenic. The phage DNA may be induced whereby it becomes disassociated from the cell DNA and enters the lytic life cycle. Temperate phage may transfer bacterial cell DNA from one cell to another to produce a recombinant.
Mosby's Dental Dictionary:

bacteriophage

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n

Any virus that causes lysis of host bacteria.

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categories related to 'bacteriophage'

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Wikipedia on Answers.com:

Bacteriophage

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The structure of a typical myovirus bacteriophage

A bacteriophage (from 'bacteria' and Greek φαγεῖν phagein "to devour") 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. The genetic material can be ssRNA, dsRNA, ssDNA, or dsDNA ('ss-' or 'ds-' prefix denotes single-strand or double-strand) along with either circular or linear arrangement.

Bacteriophages are among the most common and diverse entities in the biosphere.[1] The term is commonly used in its shortened form, phage.

Phages are widely distributed in locations populated by bacterial hosts, such as soil or the intestines of animals. One of the densest natural sources for phages and other viruses is sea water, where up to 9×108 virions per milliliter have been found in microbial mats at the surface,[2] and up to 70% of marine bacteria may be infected by phages.[3] They have been used for over 90 years as an alternative to antibiotics in the former Soviet Union and Eastern Europe as well as in France.[4] They are seen as a possible therapy against multi-drug-resistant strains of many bacteria.[5]

Contents

Classification

The dsDNA tailed phages, or Caudovirales, account for 95% of all the phages reported in the scientific literature, and possibly make up the majority of phages on the planet.[1] However, there are other phages that occur abundantly in the biosphere, phages with different virions, genomes and lifestyles. Phages are classified by the International Committee on Taxonomy of Viruses (ICTV) according to morphology and nucleic acid.

19 families are currently recognised that infect bacteria and archaea. Of these only 2 families have RNA genomes and only 5 families are enveloped. Of the viral families with DNA genomes only 2 have single stranded genomes. Eight of the viral families with DNA genomes have circular genomes while nine have linear genomes. Nine families infect bacteria only; nine infect archaea only; and one (Tectiviridae) infects both bacteria and archaea.

ICTV classification of phages[1]
Order Family Morphology Nucleic acid Examples
Caudovirales Myoviridae Non-enveloped, contractile tail Linear dsDNA T4 phage
Siphoviridae Non-enveloped, non-contractile tail (long) Linear dsDNA λ phage, Bacteriophage T5
Podoviridae Non-enveloped, non-contractile tail (short) Linear dsDNA T7 phage
Ligamenvirales Lipothrixviridae Enveloped, rod-shaped Linear dsDNA Thermoproteus tenax virus 1
Rudiviridae Non-enveloped, rod-shaped Linear dsDNA
Unassigned Ampullaviridae Enveloped, bottle-shaped Linear dsDNA
Bacilloviridae Non-enveloped, rod-shaped Linear dsDNA
Bicaudaviridae Non-enveloped, lemon-shaped Circular dsDNA
Clavaviridae Non-enveloped, rod-shaped Circular dsDNA
Corticoviridae Non-enveloped, isometric Circular dsDNA
Cystoviridae Enveloped, spherical Segmented dsRNA
Fuselloviridae Non-enveloped, lemon-shaped Circular dsDNA
Globuloviridae Enveloped, isometric Linear dsDNA
Guttavirus Non-enveloped, ovoid Circular dsDNA
Inoviridae Non-enveloped, filamentous Circular ssDNA
Leviviridae Non-enveloped, isometric Linear ssRNA
Microviridae Non-enveloped, isometric Circular ssDNA
Plasmaviridae Enveloped, pleomorphic Circular dsDNA
Tectiviridae Non-enveloped, isometric Linear dsDNA

History

Since ancient times, there have been documented reports of river waters having the ability to cure infectious diseases, such as leprosy. In 1896, Ernest Hanbury Hankin reported that something in the waters of the Ganges and Yamuna rivers in India had marked antibacterial action against cholera and could pass through a very fine porcelain filter. In 1915, British bacteriologist Frederick Twort, superintendent of the Brown Institution of London, discovered a small agent that infected and killed bacteria. He believed that the agent must be one of the following:

  1. a stage in the life cycle of the bacteria;
  2. an enzyme produced by the bacteria themselves; or
  3. a virus that grew on and destroyed the bacteria.

Twort's work was interrupted by the onset of World War I and shortage of funding.

Independently, French-Canadian microbiologist Félix d'Hérelle, working at the Pasteur Institute in Paris, announced on September 3, 1917, that he had discovered "an invisible, antagonistic microbe of the dysentery bacillus". For d’Hérelle, there was no question as to the nature of his discovery: "In a flash I had understood: what caused my clear spots was in fact an invisible microbe ... a virus parasitic on bacteria."[6] D'Hérelle called the virus a bacteriophage or bacteria-eater (from the Greek phagein meaning to eat). He also recorded a dramatic account of a man suffering from dysentery who was restored to good health by the bacteriophages.[7]

In 1923, the Eliava Institute was opened in Tbilisi, Georgia, to research this new science and put it into practice.

In 1969 Max Delbrück, Alfred Hershey and Salvador Luria were awarded the Nobel Prize in Physiology and Medicine for their discoveries of the replication of viruses and their genetic structure.

Replication

Bacteriophages may have a lytic cycle or a lysogenic cycle, and a few viruses are capable of carrying out both. With lytic phages such as the T4 phage, bacterial cells are broken open (lysed) and destroyed after immediate replication of the virion. As soon as the cell is destroyed, the phage progeny can find new hosts to infect. Lytic phages are more suitable for phage therapy. Some lytic phage undergo a phenomenon known as lysis inhibition where completed phage progeny will not immediately lyse out of the cell if there is a high extracellular phage concentrations. This mechanism is not identical to that of temperate phage going dormant and is usually temporary.

In contrast, the lysogenic cycle does not result in immediate lysing of the host cell. Those phages able to undergo lysogeny are known as temperate phages. Their viral genome will integrate with host DNA and replicate along with it fairly harmlessly, or may even become established as a plasmid. The virus remains dormant until host conditions deteriorate, perhaps due to depletion of nutrients, then the endogenous phages (known as prophages) become active. At this point they initiate the reproductive cycle, resulting in lysis of the host cell. As the lysogenic cycle allows the host cell to continue to survive and reproduce, the virus is reproduced in all of the cell’s offspring.

Sometimes prophages may provide benefits to the host bacterium while they are dormant by adding new functions to the bacterial genome in a phenomenon called lysogenic conversion. An eminent example is the conversion of a harmless strain of Vibrio cholerae by a phage into a highly virulent one, which causes cholera.

Attachment and penetration

An electron micrograph of bacteriophages attached to a bacterial cell. These viruses are the size and shape of coliphage T1

To enter a host cell, bacteriophages attach to specific receptors on the surface of bacteria, including lipopolysaccharides, teichoic acids, proteins, or even flagella. This specificity means that a bacteriophage can infect only certain bacteria bearing receptors to which they can bind, which in turn determines the phage's host range. Host growth conditions also influence the ability of the phage to attach and invade bacteria.[8] As phage virions do not move independently, they must rely on random encounters with the right receptors when in solution (blood, lymphatic circulation, irrigation, soil water, etc.).

Myovirus bacteriophages use a hypodermic syringe-like motion to inject their genetic material into the cell. After making contact with the appropriate receptor, the tail fibers flex to bring the base plate closer to the surface of the cell; this is known as reversible binding. Once attached completely, irreversible binding is initiated and the tail contracts, possibly with the help of ATP present in the tail,[3] injecting genetic material through the bacterial membrane. Podoviruses lack a elongated tail sheath similar to that of a myovirus, so they instead use their small tooth-like tail fibers to enzymatically degrade a portion of the cell membrane before inserting its genetic material.

Synthesis of proteins and nucleic acid

Within minutes, bacterial ribosomes start translating viral mRNA into protein. For RNA-based phages, RNA replicase is synthesized early in the process. Proteins modify the bacterial RNA polymerase so that it preferentially transcribes viral mRNA. The host’s normal synthesis of proteins and nucleic acids is disrupted, and it is forced to manufacture viral products instead. These products go on to become part of new virions within the cell, helper proteins that help assemble the new virions, or proteins involved in cell lysis. Walter Fiers (University of Ghent, Belgium) was the first to establish the complete nucleotide sequence of a gene (1972) and of the viral genome of Bacteriophage MS2 (1976).[9]

Virion assembly

In the case of the T4 phage, the construction of new virus particles involves the assistance of helper proteins. The base plates are assembled first, with the tails being built upon them afterwards. The head capsids, constructed separately, will spontaneously assemble with the tails. The DNA is packed efficiently within the heads. The whole process takes about 15 minutes.

Diagram of a typical tailed bacteriophage structure

Release of virions

Phages may be released via cell lysis, by extrusion, or, in a few cases, by budding. Lysis, by tailed phages, is achieved by an enzyme called endolysin, which attacks and breaks down the cell wall peptidoglycan. An altogether different phage type, the filamentous phages, make the host cell continually secrete new virus particles. Released virions are described as free, and, unless defective, are capable of infecting a new bacterium. Budding is associated with certain Mycoplasma phages. In contrast to virion release, phages displaying a lysogenic cycle do not kill the host but, rather, become long-term residents as prophage.

Genome structure

Bacteriophage genomes are especially mosaic: the genome of any one phage species appears to be composed of numerous individual modules. These modules may be found in other phage species in different arrangements. Bacteriophages with mycobacterial hosts have provided excellent examples of this mosaicism. In these mycobacteriophages, genetic assortment may be the result of repeated instances of site-specific recombination and illegitimate recombination (the result of phage genome acquisition of bacterial host genetic sequences). [10]

Phage therapy

Phages were discovered to be anti-bacterial agents and were used throughout the 1940s in the Soviet Union for treating bacterial infections. They had widespread use including treating soldiers in the Red Army. However, they were abandoned for general use in the west for several reasons:

  • Medical trials were carried out, but a basic lack of understanding of phages made these invalid.
  • Phage therapy was seen as untrustworthy, because many of the trials were conducted on totally unrelated diseases such as allergies and viral infections.
  • Antibiotics were discovered and marketed widely. They were easier to make, store and to prescribe.
  • Russian research continued but was published in Russian or Georgian, and was unavailable internationally for many years.

Their use has continued since the end of the Cold War in Georgia and elsewhere in Eastern Europe. A monograph written by Nina Chanishvili in 2009, in Tbilisi, Georgia[11] gave a thorough analysis of the results of phage therapy. It was based on the data given in the old Soviet scientific literature.

The first regulated clinical trial of efficacy in Western Europe (against ear infections caused by Pseudomonas aeruginosa) was reported in the journal Clinical Otolaryngology in August 2009.[12] Meanwhile, Western scientists are developing engineered viruses to overcome antibiotic resistance, and experimenting with tumor-suppressing agents.[13] One potential treatment currently under development is a phage designed to destroy MRSA.[14]

In the environment

Metagenomics has allowed the in-water detection of bacteriophages that was not possible previously. These investigations revealed that phage are much more abundant in the water column of both freshwater and marine habitats than previously thought[15] and that they can cause significant mortality of bacterioplankton. Methods in phage community ecology have been developed to assess phage-induced mortality of bacterioplankton and its role for food web process and biogeochemical cycle to genetically fingerprint phage communities or populations and estimate viral biodiversity by metagenomics. The lysis of bacteria by phages releases organic carbon that was previously particulate (cells) into dissolved forms, which makes the carbon more available to other organisms. Phages are not only the most abundant biological entities but probably also the most diverse ones. The majority of the sequence data obtained from phage communities has no equivalent in databases. These data and other detailed analyses indicate that phage-specific genes and ecological traits are much more frequent than previously thought. In order to reveal the meaning of this genetic and ecological versatility, studies have to be performed with communities and at spatiotemporal scales relevant for microorganisms.[1]

Bacteriophages have also been used in hydrological tracing and modelling in river systems especially where surface water and groundwater interactions occur. The use of phages is preferred to the more conventional dye marker because they are significantly less absorbed when passing through ground-waters and they are readily detected at very low concentrations.[16]

Role in food fermentation

A broad number of food products, commodity chemicals, and biotechnology products are manufactured industrially by large-scale bacterial fermentation of various organic substrates. Because enormous amounts of bacteria are being cultivated each day in large fermentation vats, the risk of bacteriophage contamination could rapidly bring fermentation to a halt. The resulting economic setback is a serious threat in these industries. The relationship between bacteriophages and their bacterial hosts is very important in the context of the food fermentation industry. Sources of phage contamination, measures to control their propagation and dissemination, and biotechnological defense strategies developed to restrain phages are of interest. The dairy fermentation industry has openly acknowledged the problem of phages and has been working with academia and starter culture companies to develop defense strategies and systems to curtail the propagation and evolution of phages for decades.[1]

Other areas of use

In August 2006, the United States Food and Drug Administration (FDA) approved LMP-102 (now ListShield) as a food additive to target and kill Listeria monocytogenes. LMP-102 was approved for treating ready-to-eat (RTE) poultry and meat products.[17] In October of that year, following the food additive approval of LMP-102 by Intralytix, the FDA approved a product by EBI using bacteriophages on cheese to kill the Listeria monocytogenes bacteria, giving them GRAS status (Generally Recognized As Safe).[18] In July 2007, the same bacteriophages were approved for use on all food products.[19] There is on going research in the field of food safety to see if lytic phage are a viable option to control other food-borne pathogens in various food products.

Government agencies in the West have for several years been looking to Georgia and the Former Soviet Union for help with exploiting phages for counteracting bioweapons and toxins, such as anthrax and botulism.[20] There are many developments with this among research groups in the US. Other uses include spray application in horticulture for protecting plants and vegetable produce from decay and the spread of bacterial disease. Other applications for bacteriophages are as a biocide for environmental surfaces, e.g., in hospitals, and as a preventative treatment for catheters and medical devices prior to use in clinical settings. The technology for phages to be applied to dry surfaces, e.g., uniforms, curtains, even sutures for surgery now exists. Clinical trials reported in the Lancet[12] show success in veterinary treatment of pet dogs with otitis.

Phage display is a different use of phages involving a library of phages with a variable peptide linked to a surface protein. Each phage's genome encodes the variant of the protein displayed on its surface (hence the name), providing a link between the peptide variant and its encoding gene. Variant phages from the library can be selected through their binding affinity to an immobilized molecule (e.g., Botulism toxin) to neutralize it. The bound selected phages can be multiplied by re-infecting a susceptible bacterial strain, thus allowing them to retrieve the peptides encoded in them for further study.[citation needed]

The SEPTIC bacterium sensing and identification method utilizes the ion emission and its dynamics during phage infection and offers high specificity and speed for detection.[citation needed]

Model bacteriophages

Following is a list of bacteriophages that are extensively studied:

Cultural references

  • In 1926 in the Pulitzer prize-winning novel Arrowsmith, Sinclair Lewis fictionalized the application of bacteriophages as a therapeutic agent.

See also

References

  1. ^ a b c d e Mc Grath S and van Sinderen D (editors). (2007). Bacteriophage: Genetics and Molecular Biology (1st ed.). Caister Academic Press. ISBN 978-1-904455-14-1. [1]. http://www.horizonpress.com/phage. 
  2. ^ Wommack, K. E.; Colwell, R. R. (2000). "Virioplankton: Viruses in Aquatic Ecosystems". Microbiology and Molecular Biology Reviews 64 (1): 69–114. doi:10.1128/MMBR.64.1.69-114.2000. PMC 98987. PMID 10704475.  edit
  3. ^ a b Prescott, L. (1993). Microbiology, Wm. C. Brown Publishers, ISBN 0-697-01372-3
  4. ^ BBC Horizon (1997): The Virus that Cures – Documentary about the history of phage medicine in Russia and the West
  5. ^ BBC article: The drugs don't work - so what will? (8 April 2011)
  6. ^ Félix d'Hérelles (1917). "Sur un microbe invisible antagoniste des bacilles dysentériques" (PDF). Comptes rendus Acad Sci Paris. 165: 373–5. Archived from the original on 2010-12-04. http://www.webcitation.org/5uicsPk41. Retrieved 2010-09-05. 
  7. ^ Félix d'Hérelle (1949). "The bacteriophage" (PDF). Science News 14: 44–59. http://mmbr.asm.org/cgi/reprint/40/4/793.pdf. Retrieved 2010-09-05. 
  8. ^ Gabashvili, I.; Khan, S.; Hayes, S.; Serwer, P. (1997). "Polymorphism of bacteriophage T7". Journal of Molecular Biology 273 (3): 658. doi:10.1006/jmbi.1997.1353. PMID 9356254.  edit
  9. ^ Fiers, W.; Contreras, R.; Duerinck, F.; Haegeman, G.; Iserentant, D.; Merregaert, J.; Min Jou, W.; Molemans, F. et al. (1976). "Complete nucleotide sequence of bacteriophage MS2 RNA: primary and secondary structure of the replicase gene". Nature 260 (5551): 500–7. Bibcode 1976Natur.260..500F. doi:10.1038/260500a0. PMID 1264203.  edit
  10. ^ Morris, P.; Marinelli, L. J., Jacobs-Sera, D., Hendrix, R. W., Hatfull, G. F. (4 January 2008). "Genomic Characterization of Mycobacteriophage Giles: Evidence for Phage Acquisition of Host DNA by Illegitimate Recombination". Journal of Bacteriology 190 (6): 2172–2182. doi:10.1128/JB.01657-07. 
  11. ^ Nina Chanishvili (2009) "A Literature Review of the Practical Application of Bacteriophage Research", p184
  12. ^ a b Wright, A.; Hawkins, C.; Anggård, E.; Harper, D. (2009). "A controlled clinical trial of a therapeutic bacteriophage preparation in chronic otitis due to antibiotic-resistant Pseudomonas aeruginosa; a preliminary report of efficacy". Clinical Otolaryngology 34 (4): 349–357. doi:10.1111/j.1749-4486.2009.01973.x. PMID 19673983.  edit
  13. ^ Breitbach, Caroline J.; Burke, James; Jonker, Derek; Stephenson, Joe; Haas, Andrew R.; Chow, Laura Q. M.; Nieva, Jorge; Hwang, Tae-Ho et al. (1 September 2011). "Intravenous delivery of a multi-mechanistic cancer-targeted oncolytic poxvirus in humans". Nature 477 (7362): 99–102. doi:10.1038/nature10358. PMID 21886163. http://www.nature.com/nature/journal/v477/n7362/full/nature10358.html. Retrieved 17 September 2011. 
  14. ^ British biotech boasts antibiotic breakthrough - Biotechnology, bacteriophages, bacteria - Australian Life Scientist
  15. ^ Breitbart, M, P Salamon, B Andresen, J Mahaffy, A Segall, D Mead, F Azam, F Rohwer (2002) Genomic analysis of uncultured marine viral communities. Proceedings of the National Academy USA. 99:14250-14255.
  16. ^ Martin, C. (1988). "The Application of Bacteriophage Tracer Techniques in South West Water". Water and Environment Journal 2: 638. doi:10.1111/j.1747-6593.1988.tb01352.x.  edit
  17. ^ govpulse | Food Additives Permitted for Direct Addition to Food for Human Consumption; Bacteriophage Preparation
  18. ^ U.S. FDA/CFSAN: Agency Response Letter: GRAS Notice No. GRN 000198[dead link]
  19. ^ U.S. FDA/CFSAN: Agency Response Letter: GRAS Notice No. GRN 000218[dead link]
  20. ^ The New York Times: Studying anthrax in a Soviet-era lab - with Western funding
  21. ^ Agreement: Use of the Esther M. Zimmer Lederberg Memorial Web Site

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phage
transduction
bactericide

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