Virulent Bacteriophages - Lytic Cycle (cytoplasmic Viral Replication) ~ Do News

...of bacteriophage To discuss the infectious process and the lytic multiplication cycle To explain the lysogenic cycle and its regulation. In the more complex phages like T4 the tail is surrounded by a contractile sheath which contracts during infection of the bacterium. At the end of the tail the more...Endolysins are the lytic enzymes used by phages at the end of the replication cycle to degrade bacterial peptidoglycan (PG) from within, resulting in a 2010). Another multi-threaded mechanism protecting bacteria against lytic and temperate phage infection is the bacteriophage exclusion...Bacteriophages. Bacteria-infecting viruses. Bacteriophages. This is the currently selected item.In the lytic cycle, the genetic material of virus transcribes and produces capsid proteins by using host molecular machinery. Then the capsid protein joins around the viral genetic material and new phage particles are produced. The lytic cycle ends when these new phage particles released out of the cell...The multiplication cycle of the T-even bacteriophage ends with the lysis of the host cell where the term "Lytic This article will only discuss the lytic cycle using the T-even bacteriophage infecting its host, E. coli, as an A few minutes after infection, complete phages cannot be found in the host cell.

PDF Applications of bacteriophages versus phage enzymes to combat and...

2.4 Replication cycles of lytic and lysogenic bacteriophages. 11. 2.5 Resistance pattern in Shigella During infection, it undergoes conformational change and allows the penetration of DNA from the viral core The excised DNA will enter the lytic cycle that causes death in bacterial cell (Todar 2012). Increase of phage predation will reduce the bacterial density, which ultimately ends the outbreak by...The structure of a typical myovirus bacteriophage. Anatomy and infection cycle of phage T4. An example of a bacteriophage known to follow the lysogenic cycle and the lytic cycle is the phage For instance, infection of Pseudomonas aeruginosa by the temperate phage PaP3 changed the...In lysogenic cycle, the lytic (vegetative) phage becomes integrated with the host cell chromosomes and is converted into prophage without lysis of bacterial cell. Bacteriophage infection in S. typhi confers a new antigenic surface structure on the host.Lytic infection ends in cell-death (the membrane ruptures). Lysogenic infection means that the virus "hibernates" inside the cell for a time and can bud The cash cycle starts when you pay your supplier and ends when your buyer pays you. The operating cycle starts with acquiring of inventory or raw...

Bacteria-infecting viruses. The lytic and lysogenic cycles.

Lytic cycle is one one of the two alternative life cycles of a virus inside a host cell, whereby the virus that has entered a cell takes over the cell's replication mechanism, makes viral DNA and viral proteins, and then lyses (breaks open) the cell...Bacteriophage and bacteriophage typing A bacteriophage, or phage, is a virus that infects a bacterial cell At the end of the lytic cycle, the phage directs the host cell to produce the enzyme, lysozyme, that See also Bacteria and bacterial infection; Biotechnology; Cell cycle (prokaryotic)...The lytic cycle of bacteriophage infection ends with the _____. A plant that has been raised in a sterile environment shows symptoms of a viral infection. How would you explain this? The viral infection was acquired by vertical transmission.Bacteriophages are diverse group of viruses which are easily manipulated and therefore they have potential uses in biotechnology, research, and therapeutics. The aim of this review article is to enable the wide range of researchers, scientists, and biotechnologist who are putting phages into practice, to...Video of The cycle of infection results in the death of the host cell and the release of many virus particles, called virions. Interestingly, the lytic cycle does not always happen immediately. Sometimes, rather than producing virions, phage nucleic acid incorporates in the host cell DNA.

Jump to navigation Jump to go looking "Phage" redirects right here. For different makes use of, see Phage (disambiguation).

Atomic structural fashion of bacteriophage T4 The structure of a regular myovirus bacteriophage Anatomy and infection cycle of phage T4.

A bacteriophage (/bækˈtɪərioʊfeɪdʒ/), additionally identified informally as a phage (/ˈfeɪdʒ/), is an endemic that infects and replicates inside bacteria and archaea. The time period was derived from "bacteria" and the Greek φαγεῖν (phagein), that means "to devour". Bacteriophages are composed of proteins that encapsulate a DNA or RNA genome, and will have structures which might be either easy or elaborate. Their genomes would possibly encode as few as 4 genes (e.g. MS2) and as many as masses of genes. Phages mirror within the bacterium following the injection of their genome into its cytoplasm.

Bacteriophages are among the maximum common and numerous entities in the biosphere.[1] Bacteriophages are ubiquitous viruses, found anyplace bacteria exist. It is estimated there are greater than 1031 bacteriophages on the planet, greater than each different organism on Earth, together with micro organism, mixed.[2] Viruses are the maximum considerable organic entity in the water column of the global's oceans, and the second greatest part of biomass after prokaryotes,[3] the place as much as 9x108virions consistent with millilitre had been present in microbial mats at the floor,[4] and as much as 70% of marine bacteria is also infected by means of phages.[5]

Phages had been used since the past due twentieth century as a substitute for antibiotics in the former Soviet Union and Central Europe, in addition to in France.[6][7] They are seen as a possible remedy in opposition to multi-drug-resistant traces of many micro organism (see phage treatment).[8] On the different hand, phages of Inoviridae have been proven to complicate biofilms interested by pneumonia and cystic fibrosis and to refuge the micro organism from drugs supposed to eradicate illness, thus selling chronic infection.[9]

Classification

Bacteriophages happen abundantly in the biosphere, with different genomes, and life. Phages are classified by way of the International Committee on Taxonomy of Viruses (ICTV) in step with morphology and nucleic acid.

Bacteriophage P22, a member of the Podoviridae through morphology because of its quick, non-contractile tail ICTV classification of prokaryotic (bacterial and archaeal) viruses[1] Order Family Morphology Nucleic acid Examples Belfryvirales Turriviridae Enveloped, isometric Linear dsDNA Caudovirales Ackermannviridae Nonenveloped, contractile tail Linear dsDNA Myoviridae Nonenveloped, contractile tail Linear dsDNA T4, Mu, P1, P2 Siphoviridae Nonenveloped, noncontractile tail (long) Linear dsDNA λ, T5, HK97, N15 Podoviridae Nonenveloped, noncontractile tail (quick) Linear dsDNA T7, T3, Φ29, P22 Halopanivirales Sphaerolipoviridae Enveloped, isometric Linear dsDNA Haloruvirales Pleolipoviridae Enveloped, pleomorphic Circular ssDNA, circular dsDNA, or linear dsDNA Kalamavirales Tectiviridae Nonenveloped, isometric Linear dsDNA Levivirales Leviviridae Nonenveloped, isometric Linear ssRNA MS2, Qβ Ligamenvirales Lipothrixviridae Enveloped, rod-shaped Linear dsDNA Acidianus filamentous virus 1 Rudiviridae Nonenveloped, rod-shaped Linear dsDNA Sulfolobus islandicus rod-shaped virus 1 Mindivirales Cystoviridae Enveloped, spherical Segmented dsRNA Φ6 Petitvirales Microviridae Nonenveloped, isometric Circular ssDNA ΦX174 Tubulavirales Inoviridae Nonenveloped, filamentous Circular ssDNA M13 Vinavirales Corticoviridae Nonenveloped, isometric Circular dsDNA PM2 Unassigned Ampullaviridae Enveloped, bottle-shaped Linear dsDNA Bicaudaviridae Nonenveloped, lemon-shaped Circular dsDNA Clavaviridae Nonenveloped, rod-shaped Circular dsDNA Finnlakeviridae dsDNA FLiP[10]Fuselloviridae Nonenveloped, lemon-shaped Circular dsDNA Globuloviridae Enveloped, isometric Linear dsDNA Guttaviridae Nonenveloped, ovoid Circular dsDNA Plasmaviridae Enveloped, pleomorphic Circular dsDNA Portogloboviridae Enveloped, isometric Circular dsDNA Spiraviridae Nonnveloped, rod-shaped Circular ssDNA Tristromaviridae Enveloped, rod-shaped Linear dsDNA

It has been advised that individuals of Picobirnaviridae infect bacteria, however now not mammals.[11]

Another proposed family is "Autolykiviridae" (dsDNA).[12]

History

Félix d'Herelle

In 1896, Ernest Hanbury Hankin reported that something in the waters of the Ganges and Yamuna rivers in India had a marked antibacterial motion against cholera and it might cross through a very superb porcelain filter.[13] In 1915, British bacteriologist Frederick Twort, superintendent of the Brown Institution of London, came upon a small agent that inflamed and killed bacteria. He believed the agent should be one of the following:

a degree in the lifestyles cycle of the bacteria an enzyme produced by means of the micro organism themselves, or an endemic that grew on and destroyed the micro organism[14]

Twort's analysis used to be interrupted by way of the onset of World War I, as well as a scarcity of investment and the discoveries of antibiotics.

Independently, French-Canadian microbiologist Félix d'Hérelle, working at the Pasteur Institute in Paris, introduced on 3 September 1917, that he had came upon "an invisible, antagonistic microbe of the dysentery bacillus". For d'Hérelle, there was once 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."[15] D'Hérelle called the virus a bacteriophage, a bacteria-eater (from the Greek phagein meaning "to devour"). He also recorded a dramatic account of a man suffering from dysentery who was once restored to excellent well being by the bacteriophages.[16] It was D'Herelle who carried out a lot research into bacteriophages and introduced the concept of phage treatment.[17]

More than a part a century later, in 1969, Max Delbrück, Alfred Hershey, and Salvador Luria have been awarded the Nobel Prize in Physiology or Medicine for his or her discoveries of the replication of viruses and their genetic construction.[18]

Uses

Phage remedy Main article: Phage therapy

Phages were found out to be antibacterial agents and were utilized in the former Soviet Republic of Georgia (pioneered there by means of Giorgi Eliava with assist from the co-discoverer of bacteriophages, Félix d'Herelle) all the way through the Nineteen Twenties and 1930s for treating bacterial infections. They had standard use, including treatment of soldiers in the Red Army. However, they have been abandoned for general use in the West for a number of causes:

Antibiotics have been discovered and advertised extensively. They were easier to make, retailer, and to prescribe. Medical trials of phages have been performed, however a elementary lack of working out raised questions about the validity of these trials.[19] Publication of analysis in the Soviet Union was mainly in the Russian or Georgian languages and for many years, used to be not adopted internationally.

The use of phages has persevered since the finish of the Cold War in Russia,[20] Georgia and in different places in Central and Eastern Europe. The first regulated, randomized, double-blind clinical trial used to be reported in the Journal of Wound Care in June 2009, which evaluated the protection and efficacy of a bacteriophage cocktail to regard infected venous ulcers of the leg in human patients.[21] The FDA authorized the find out about as a Phase I medical trial. The learn about's results demonstrated the protection of healing application of bacteriophages, however didn't display efficacy. The authors defined that the use of positive chemical compounds which are section of standard wound care (e.g. lactoferrin or silver) will have interfered with bacteriophage viability.[21] Shortly after that, any other controlled clinical trial in Western Europe (treatment of ear infections caused by Pseudomonas aeruginosa) used to be reported in the journal Clinical Otolaryngology in August 2009.[22] The study concludes that bacteriophage arrangements had been protected and efficient for treatment of persistent ear infections in humans. Additionally, there were numerous animal and other experimental medical trials evaluating the efficacy of bacteriophages for various diseases, reminiscent of infected burns and wounds, and cystic fibrosis associated lung infections, among others.[22]

Meanwhile, bacteriophage researchers have been growing engineered viruses to overcome antibiotic resistance, and engineering the phage genes liable for coding enzymes that degrade the biofilm matrix, phage structural proteins, and the enzymes responsible for lysis of the bacterial cell wall.[4][5][6] There were results showing that T4 phages which are small in size and short-tailed, will also be useful in detecting E.coli in the human body.[23]

Therapeutic efficacy of a phage cocktail used to be evaluated in a mice type with nasal infection of multidrug-resistant (MDR) A. baumannii. Mice handled with the phage cocktail showed a 2.3-fold higher survival charge than the ones untreated in seven days submit infection.[24] In 2017 a affected person with a pancreas compromised by way of MDR A. baumannii was once placed on several antibiotics, regardless of this the affected person's well being endured to go to pot during a four-month period. Without efficient antibiotics the affected person used to be subjected to phage treatment the use of a phage cocktail containing nine different phages that had been demonstrated to be efficient in opposition to MDR A. baumannii. Once on this treatment the patient's downward scientific trajectory reversed, and returned to well being.[25]

D'Herelle "quickly learned that bacteriophages are found wherever bacteria thrive: in sewers, in rivers that catch waste runoff from pipes, and in the stools of convalescent patients."[26] This comprises rivers historically thought to have therapeutic powers, including India's Ganges River.[27]

Other

Food business – Since 2006, the United States Food and Drug Administration (FDA) and United States Department of Agriculture (USDA) have approved several bacteriophage products. LMP-102 (Intralytix) used to be approved for treating ready-to-eat (RTE) poultry and meat products. In that same yr, the FDA approved LISTEX (evolved and produced through Micreos) the use of bacteriophages on cheese to kill Listeria monocytogenes bacteria, with the intention to give them generally recognized as protected (GRAS) standing.[28] In July 2007, the identical bacteriophage had been approved for use on all meals products.[29] In 2011 USDA showed that LISTEX is a clean label processing assist and is incorporated in USDA.[30] Research in the field of meals protection is continuous to see if lytic phages are a viable option to regulate different food-borne pathogens in quite a lot of food merchandise.

Dairy industry – Bacteriophages found in the atmosphere can cause fermentation failures of cheese starter cultures. In order to steer clear of this, mixed-strain starter cultures and tradition rotation regimes can be utilized.[31]

Diagnostics – In 2011, the FDA cleared the first bacteriophage-based product for in vitro diagnostic use.[32] The KeyPath MRSA/MSSA Blood Culture Test makes use of a cocktail of bacteriophage to detect Staphylococcus aureus in certain blood cultures and determine methicillin resistance or susceptibility. The take a look at returns leads to about five hours, compared to two to a few days for usual microbial identification and susceptibility take a look at methods. It used to be the first accelerated antibiotic-susceptibility take a look at approved by means of the FDA.[33]

Counteracting bioweapons and toxins – Government agencies in the West have for several years been looking to Georgia and the former Soviet Union for assist with exploiting phages for counteracting bioweapons and toxins, similar to anthrax and botulism.[34] Developments are proceeding among analysis teams in the U.S. Other makes use of include spray application in horticulture for protecting crops and vegetable produce from decay and the unfold of bacterial disease. Other applications for bacteriophages are as biocides for environmental surfaces, e.g., in hospitals, and as preventative therapies for catheters and clinical gadgets before use in clinical settings. The technology for phages to be carried out to dry surfaces, e.g., uniforms, curtains, or even sutures for surgery now exists. Clinical trials reported in Clinical Otolaryngology[22] display success in veterinary remedy of pet canine with otitis.

The SEPTIC bacterium sensing and identification manner makes use of the ion emission and its dynamics all over phage infection and gives top specificity and speed for detection.[35]

Phage show is a unique use of phages involving a library of phages with a variable peptide connected to a floor protein. Each phage genome encodes the variant of the protein displayed on its floor (therefore the name), providing a link between the peptide variant and its encoding gene. Variant phages from the library could also be decided on through their binding affinity to an immobilized molecule (e.g., botulism toxin) to neutralize it. The sure, decided on phages will also be multiplied by means of reinfecting a vulnerable bacterial pressure, thus allowing them to retrieve the peptides encoded in them for additional study.[36]

Antimicrobial drug discovery – Phage proteins steadily have antimicrobial activity and would possibly function leads for peptidomimetics, i.e. drugs that mimic peptides.[37]Phage-ligand era makes use of phage proteins for various programs, such as binding of bacteria and bacterial components (e.g. endotoxin) and lysis of micro organism.[38]

Basic analysis – Bacteriophages are essential model organisms for learning rules of evolution and ecology.[39]

Replication

Diagram of the DNA injection procedure

Bacteriophages can have a lytic cycle or a lysogenic cycle. With lytic phages reminiscent of the T4 phage, bacterial cells are broken open (lysed) and destroyed after fast replication of the virion. As soon as the cell is destroyed, the phage progeny can in finding new hosts to contaminate. Lytic phages are more suitable for phage treatment. Some lytic phages undergo a phenomenon referred to as lysis inhibition, where finished phage progeny is not going to immediately lyse out of the mobile if extracellular phage concentrations are top. This mechanism is not identical to that of temperate phage going dormant and normally, is brief.

In contrast, the lysogenic cycle does not result in rapid lysing of the host cellular. Those phages ready to undergo lysogeny are referred to as temperate phages. Their viral genome will integrate with host DNA and reflect alongside with it, fairly harmlessly, or may even change into established as a plasmid. The virus stays dormant until host stipulations deteriorate, possibly because of depletion of vitamins, then, the endogenous phages (known as prophages) grow to be energetic. At this level they start up the reproductive cycle, resulting in lysis of the host cellular. As the lysogenic cycle allows the host cell to continue to live to tell the tale and reproduce, the virus is replicated in all offspring of the cell. An instance of a bacteriophage identified to observe the lysogenic cycle and the lytic cycle is the phage lambda of E. coli.[40]

Sometimes prophages would possibly supply benefits to the host bacterium while they're dormant by way of including new functions to the bacterial genome, in a phenomenon referred to as lysogenic conversion. Examples are the conversion of harmless lines of Corynebacterium diphtheriae or Vibrio cholerae by bacteriophages, to highly virulent ones that purpose diphtheria or cholera, respectively.[41][42] Strategies to battle sure bacterial infections by means of focused on these toxin-encoding prophages were proposed.[43]

Attachment and penetration In this electron micrograph of bacteriophages hooked up to a bacterial cellular, the viruses are the length and shape of coliphage T1

Bacterial cells are safe via a cellular wall of polysaccharides, which can be necessary virulence components protective bacterial cells towards each immune host defenses and antibiotics.[44] To enter a host cell, bacteriophages bind to specific receptors on the surface of bacteria, together with lipopolysaccharides, teichoic acids, proteins, or even flagella. This specificity manner a bacteriophage can infect best positive micro organism bearing receptors to which they are able to bind, which in flip, determines the phage's host vary. Polysaccharide-degrading enzymes, like endolysins are virion-associated proteins to enzymatically degrade the capsular outer layer of their hosts, at the initial step of a tightly programmed phage infection process. Host enlargement conditions additionally influence the ability of the phage to attach and invade them.[45] As phage virions do not move independently, they must depend on random encounters with the proper receptors when in answer, such as blood, lymphatic stream, irrigation, soil water, and so forth.

Myovirus bacteriophages use a hypodermic syringe-like movement to inject their genetic subject matter into the cell. After contacting the appropriate receptor, the tail fibers flex to convey the base plate nearer to the surface of the mobile. This is referred to as reversible binding. Once hooked up utterly, irreversible binding is initiated and the tail contracts, possibly with the help of ATP, found in the tail,[5] injecting genetic subject matter via the bacterial membrane.[46] The injection is accomplished thru a sort of bending movement in the shaft by means of going to the facet, contracting closer to the mobile and pushing back up. Podoviruses lack an elongated tail sheath like that of a myovirus, so as a substitute, they use their small, tooth-like tail fibers enzymatically to degrade a portion of the cell membrane sooner than putting their genetic material.

Synthesis of proteins and nucleic acid

Within minutes, bacterial ribosomes get started translating viral mRNA into protein. For RNA-based phages, RNA replicase is synthesized early in the procedure. Proteins regulate the bacterial RNA polymerase so it preferentially transcribes viral mRNA. The host's customary synthesis of proteins and nucleic acids is disrupted, and it's pressured to manufacture viral products as a substitute. These merchandise cross directly to turn into part of new virions inside of the cell, helper proteins that contribute to the assemblage of new virions, or proteins occupied with cell lysis. In 1972, Walter Fiers (University of Ghent, Belgium) was once the first to establish the complete nucleotide series of a gene and in 1976, of the viral genome of bacteriophage MS2.[47] Some dsDNA bacteriophages encode ribosomal proteins, which might be thought to modulate protein translation all over phage infection.[48]

Virion meeting

In the case of the T4 phage, the construction of new virus particles comes to the help of helper proteins that act catalytically right through phage morphogenesis.[49] The base plates are assembled first, with the tails being constructed upon them in a while. The head capsids, built one after the other, will spontaneously bring together with the tails. During meeting of the phage T4 virion, the morphogenetic proteins encoded by the phage genes interact with each and every other in a characteristic series. Maintaining an appropriate stability in the quantities of each and every of those proteins produced all over viral infection appears to be vital for standard phage T4 morphogenesis.[50] The DNA is packed successfully within the heads. The whole procedure takes about quarter-hour.

Release of virions

Phages could also be released by way of cellular lysis, by extrusion, or, in a few instances, by means of budding. Lysis, by tailed phages, is achieved via an enzyme known as endolysin, which attacks and breaks down the cellular wall peptidoglycan. An altogether other phage sort, the filamentous phage, make the host mobile regularly secrete new virus debris. Released virions are described as loose, and, until faulty, are succesful of infecting a brand new bacterium. Budding is associated with positive Mycoplasma phages. In contrast to virion free up, phages exhibiting a lysogenic cycle don't kill the host however, somewhat, grow to be long-term citizens as prophage.

Communication

Research in 2017 revealed that the bacteriophage Φ3T makes a short viral protein that alerts different bacteriophages to lie dormant as a substitute of killing the host bacterium. Arbitrium is the identify given to this protein by way of the researchers who came upon it.[51][52]

Genome construction

Given the tens of millions of other phages in the setting, phage genomes are available a variety of paperwork and sizes. RNA phage reminiscent of MS2 have the smallest genomes, of only some kilobases. However, some DNA phage such as T4 will have massive genomes with loads of genes; the size and form of the capsid varies alongside with the length of the genome.[53] The biggest bacteriophage genomes succeed in a size of 735 kb.[54]

Bacteriophage genomes can be highly mosaic, i.e. the genome of many phage species appear to be composed of a lot of individual modules. These modules is also found in other phage species in different preparations. Mycobacteriophages, bacteriophages with mycobacterial hosts, have supplied superb examples of this mosaicism. In those mycobacteriophages, genetic collection may be the result of repeated cases of site-specific recombination and illegitimate recombination (the end result of phage genome acquisition of bacterial host genetic sequences).[55] Evolutionary mechanisms shaping the genomes of bacterial viruses vary between different families and rely on the kind of the nucleic acid, traits of the virion structure, in addition to the mode of the viral existence cycle.[56]

Systems biology

The box of techniques biology investigates the advanced networks of interactions within an organism, usually the usage of computational equipment and modeling.[57] For example, a phage genome that enters right into a bacterial host cell may specific loads of phage proteins which can impact the expression of a large number of host gene or the host's metabolism. All of those complex interactions may also be described and simulated in pc models.[57]

For example, infection of Pseudomonas aeruginosa by means of the temperate phage PaP3 modified the expression of 38% (2160/5633) of its host's genes. Many of those results are probably indirect, hence the problem becomes to spot the direct interactions amongst micro organism and phage.[58]

Several makes an attempt were made to map protein–protein interactions among phage and their host. For instance, bacteriophage lambda was once discovered to have interaction with its host, E. coli, by dozens of interactions. Again, the importance of many of those interactions remains unclear, however those research recommend that there perhaps are a number of key interactions and many oblique interactions whose role remains uncharacterized.[59]

In the surroundings

Main article: Marine bacteriophage

Metagenomics has allowed the in-water detection of bacteriophages that used to be not possible in the past.[60]

Also, bacteriophages had been utilized in hydrological tracing and modelling in river methods, especially the place floor water and groundwater interactions happen. The use of phages is most popular to the more conventional dye marker because they're considerably much less absorbed when passing thru flooring waters and they are readily detected at very low concentrations.[61] Non-polluted water might contain approximately 2×108 bacteriophages according to ml.[62]

Bacteriophages are thought to contribute widely to horizontal gene switch in natural environments, basically by the use of transduction, but in addition by way of transformation.[63]Metagenomics-based research also have revealed that viromes from a variety of environments harbor antibiotic-resistance genes, together with those who may just confer multidrug resistance.[64]

Model bacteriophages

The following bacteriophages are widely studied:

186 phage λ phage Φ6 phage Φ29 phage ΦX174 G4 phage M13 phage MS2 phage (23–28 nm in length)[65] N4 phage P1 phage P2 phage P4 phage R17 phage T2 phage T4 phage (169 kbp genome,[66] 2 hundred nm long[67]) T7 phage T12 phage

References

^ a b .mw-parser-output cite.quotationfont-style:inherit.mw-parser-output .quotation qquotes:"\"""\"""'""'".mw-parser-output .id-lock-free a,.mw-parser-output .citation .cs1-lock-free abackground:linear-gradient(clear,transparent),url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")appropriate 0.1em center/9px no-repeat.mw-parser-output .id-lock-limited a,.mw-parser-output .id-lock-registration a,.mw-parser-output .quotation .cs1-lock-limited a,.mw-parser-output .quotation .cs1-lock-registration abackground:linear-gradient(transparent,clear),url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")appropriate 0.1em middle/9px no-repeat.mw-parser-output .id-lock-subscription a,.mw-parser-output .quotation .cs1-lock-subscription abackground:linear-gradient(clear,clear),url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")appropriate 0.1em heart/9px no-repeat.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registrationcolor:#555.mw-parser-output .cs1-subscription span,.mw-parser-output .cs1-registration spanborder-bottom:1px dotted;cursor:assist.mw-parser-output .cs1-ws-icon abackground:linear-gradient(clear,clear),url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat.mw-parser-output code.cs1-codecolour:inherit;background:inherit;border:none;padding:inherit.mw-parser-output .cs1-hidden-errordisplay:none;font-size:100%.mw-parser-output .cs1-visible-errorfont-size:100%.mw-parser-output .cs1-maintshow:none;colour:#33aa33;margin-left:0.3em.mw-parser-output .cs1-formatfont-size:95%.mw-parser-output .cs1-kern-left,.mw-parser-output .cs1-kern-wl-leftpadding-left:0.2em.mw-parser-output .cs1-kern-right,.mw-parser-output .cs1-kern-wl-rightpadding-right:0.2em.mw-parser-output .quotation .mw-selflinkfont-weight:inheritMcGrath S and van Sinderen D (editors). 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