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99-424: Ranavirus is a genus of viruses in the family Iridoviridae . There are six other genera of viruses within the family Iridoviridae , but Ranavirus is the only one that includes viruses that are infectious to amphibians and reptiles. Additionally, it is one of the three genera within this family which infect teleost fishes , along with Lymphocystivirus and Megalocytivirus . The Ranaviruses, like

198-425: A virion , consists of nucleic acid surrounded by a protective coat of protein called a capsid . These are formed from protein subunits called capsomeres . Viruses can have a lipid "envelope" derived from the host cell membrane . The capsid is made from proteins encoded by the viral genome and its shape serves as the basis for morphological distinction. Virally-coded protein subunits will self-assemble to form

297-467: A DSB. The process starts with the key end resection factor CtlP, which mediates the end resection of DSBs, resulting in the formation of a 3' ssDNA extension. Meditated by RAD52 , the flanking homologous sequences are annealed, and forms a synapse intermediate. Then, the nonhomologous 3’ extension is removed by the ERCC1 - XPF complex through endonucleolytic cleavage, with RAD52 increasing the efficiency of

396-622: A basic optical microscope. In 2013, the Pandoravirus genus was discovered in Chile and Australia, and has genomes about twice as large as Megavirus and Mimivirus. All giant viruses have dsDNA genomes and they are classified into several families: Mimiviridae , Pithoviridae, Pandoraviridae , Phycodnaviridae , and the Mollivirus genus. Some viruses that infect Archaea have complex structures unrelated to any other form of virus, with

495-502: A bridge and bring both ends of the damaged DNA strands together. This is then followed by the processing of any non-ligatable DNA termini by a group of proteins including Artemis , PNKP , APLF and Ku, before the XRCC4 and DNA Ligase IV ligate the bridged DNA. Microhomology-mediated end joining (MMEJ), also known as alt-non-homologous end joining, is another pathway to repair DSBs. The process of MMEJ can be summarized in five steps:

594-418: A capsid diameter of 400 nm. Protein filaments measuring 100 nm project from the surface. The capsid appears hexagonal under an electron microscope, therefore the capsid is probably icosahedral. In 2011, researchers discovered the largest then known virus in samples of water collected from the ocean floor off the coast of Las Cruces, Chile. Provisionally named Megavirus chilensis , it can be seen with

693-581: A capsid, in general requiring the presence of the virus genome. Complex viruses code for proteins that assist in the construction of their capsid. Proteins associated with nucleic acid are known as nucleoproteins , and the association of viral capsid proteins with viral nucleic acid is called a nucleocapsid. The capsid and entire virus structure can be mechanically (physically) probed through atomic force microscopy . In general, there are five main morphological virus types: The poxviruses are large, complex viruses that have an unusual morphology. The viral genome

792-596: A cell, viruses exist in the form of independent viral particles, or virions , consisting of (i) genetic material , i.e., long molecules of DNA or RNA that encode the structure of the proteins by which the virus acts; (ii) a protein coat, the capsid , which surrounds and protects the genetic material; and in some cases (iii) an outside envelope of lipids . The shapes of these virus particles range from simple helical and icosahedral forms to more complex structures. Most virus species have virions too small to be seen with an optical microscope and are one-hundredth

891-639: A cellular structure, which is often seen as the basic unit of life. Viruses do not have their own metabolism and require a host cell to make new products. They therefore cannot naturally reproduce outside a host cell —although some bacteria such as rickettsia and chlamydia are considered living organisms despite the same limitation. Accepted forms of life use cell division to reproduce, whereas viruses spontaneously assemble within cells. They differ from autonomous growth of crystals as they inherit genetic mutations while being subject to natural selection. Virus self-assembly within host cells has implications for

990-499: A contributing factor in the global decline of amphibian populations. The impact of Ranaviruses on amphibian populations has been compared to the chytrid fungus Batrachochytrium dendrobatidis , the causative agent of chytridiomycosis . In the UK, the severity of disease outbreaks is thought to have increased due to climate change. Rana is derived from the Latin for "frog", reflecting

1089-493: A different DNA (or RNA) molecule. This can occur when viruses infect cells simultaneously and studies of viral evolution have shown that recombination has been rampant in the species studied. Recombination is common to both RNA and DNA viruses. Coronaviruses have a single-strand positive-sense RNA genome. Replication of the genome is catalyzed by an RNA-dependent RNA polymerase . The mechanism of recombination used by coronaviruses likely involves template switching by

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1188-648: A few hours post infection. There are several hypotheses for seasonal outbreak patterns observed for Ranavirosis mortality events. Ranaviruses grow in vitro between 8-30 °C, however for most isolates, warmer temperature result in faster viral replication. A combination of this optimal growth temperature along with shifts in larval amphibian susceptibility result in seasonal outbreak events most often observed during warm summer months. Amphibian mortality events are often observed as larval amphibians reach late Gosner stages approaching metamorphosis . As larval amphibians reach metamorphic stages of development, their immune system

1287-559: A few species, or broad for viruses capable of infecting many. Viral infections in animals provoke an immune response that usually eliminates the infecting virus. Immune responses can also be produced by vaccines , which confer an artificially acquired immunity to the specific viral infection. Some viruses, including those that cause HIV/AIDS , HPV infection , and viral hepatitis , evade these immune responses and result in chronic infections. Several classes of antiviral drugs have been developed. The English word "virus" comes from

1386-549: A fluid, by Wendell Meredith Stanley , and the invention of the electron microscope in 1931 allowed their complex structures to be visualised. Scientific opinions differ on whether viruses are a form of life or organic structures that interact with living organisms. They have been described as "organisms at the edge of life", since they resemble organisms in that they possess genes , evolve by natural selection , and reproduce by creating multiple copies of themselves through self-assembly. Although they have genes, they do not have

1485-445: A genome size of only two kilobases; the largest—the pandoraviruses —have genome sizes of around two megabases which code for about 2500 proteins. Virus genes rarely have introns and often are arranged in the genome so that they overlap . In general, RNA viruses have smaller genome sizes than DNA viruses because of a higher error-rate when replicating, and have a maximum upper size limit. Beyond this, errors when replicating render

1584-515: A host organism. Amphibian mass mortality events due to Ranavirus have been reported in Asia, Europe, North America, and South America. Ranaviruses have been isolated from wild populations of amphibians in Australia, but have not been associated with mass mortality on that continent. Synthesis of viral proteins begins within hours of viral entry with necrosis or apoptosis occurring as early as

1683-513: A ladder. The virus particles of some virus families, such as those belonging to the Hepadnaviridae , contain a genome that is partially double-stranded and partially single-stranded. For most viruses with RNA genomes and some with single-stranded DNA (ssDNA) genomes, the single strands are said to be either positive-sense (called the 'plus-strand') or negative-sense (called the 'minus-strand'), depending on if they are complementary to

1782-451: A large single linear dsDNA genome of roughly 105 kbp which codes for around 100 gene products. The main structural component of the protein capsid is the major capsid protein (MCP). Ranaviral replication is well studied using Frog virus 3 (FV3). Replication of FV3 occurs between 12 and 32 degrees Celsius. Ranaviruses enter the host cell by receptor-mediated endocytosis . Viral particles are uncoated and subsequently move into

1881-436: A life form, because they carry genetic material, reproduce, and evolve through natural selection , although they lack some key characteristics, such as cell structure, that are generally considered necessary criteria for defining life. Because they possess some but not all such qualities, viruses have been described as "organisms at the edge of life" and as replicators . Viruses spread in many ways. One transmission pathway

1980-472: A limited range of hosts and many are species-specific. Some, such as smallpox virus for example, can infect only one species—in this case humans, and are said to have a narrow host range . Other viruses, such as rabies virus, can infect different species of mammals and are said to have a broad range. The viruses that infect plants are harmless to animals, and most viruses that infect other animals are harmless to humans. The host range of some bacteriophages

2079-408: A prime target for natural selection. Segmented genomes confer evolutionary advantages; different strains of a virus with a segmented genome can shuffle and combine genes and produce progeny viruses (or offspring) that have unique characteristics. This is called reassortment or 'viral sex'. Genetic recombination is a process by which a strand of DNA (or RNA) is broken and then joined to the end of

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2178-412: A single viral particle that is released from the cell and is capable of infecting other cells of the same type. Viruses are found wherever there is life and have probably existed since living cells first evolved . The origin of viruses is unclear because they do not form fossils, so molecular techniques are used to infer how they arose. In addition, viral genetic material occasionally integrates into

2277-460: A small part of the total diversity of viruses has been studied. As of 2022, 6 realms, 10 kingdoms, 17 phyla, 2 subphyla, 40 classes, 72 orders, 8 suborders, 264 families, 182 subfamilies , 2,818 genera, 84 subgenera , and 11,273 species of viruses have been defined by the ICTV. The general taxonomic structure of taxon ranges and the suffixes used in taxonomic names are shown hereafter. As of 2022,

2376-605: A wide diversity of sizes and shapes, called ' morphologies '. In general, viruses are much smaller than bacteria and more than a thousand bacteriophage viruses would fit inside an Escherichia coli bacterium's cell. Many viruses that have been studied are spherical and have a diameter between 20 and 300 nanometres . Some filoviruses , which are filaments, have a total length of up to 1400 nm; their diameters are only about 80 nm. Most viruses cannot be seen with an optical microscope , so scanning and transmission electron microscopes are used to visualise them. To increase

2475-553: A wide variety of unusual shapes, ranging from spindle-shaped structures to viruses that resemble hooked rods, teardrops or even bottles. Other archaeal viruses resemble the tailed bacteriophages, and can have multiple tail structures. An enormous variety of genomic structures can be seen among viral species ; as a group, they contain more structural genomic diversity than plants, animals, archaea, or bacteria. There are millions of different types of viruses, although fewer than 7,000 types have been described in detail. As of January 2021,

2574-416: Is a feature of many bacterial and some animal viruses. Some viruses undergo a lysogenic cycle where the viral genome is incorporated by genetic recombination into a specific place in the host's chromosome. The viral genome is then known as a " provirus " or, in the case of bacteriophages a " prophage ". Whenever the host divides, the viral genome is also replicated. The viral genome is mostly silent within

2673-405: Is a major change in the genome of the virus. This can be a result of recombination or reassortment . The Influenza A virus is highly prone to reassortment; occasionally this has resulted in novel strains which have caused pandemics . RNA viruses often exist as quasispecies or swarms of viruses of the same species but with slightly different genome nucleoside sequences. Such quasispecies are

2772-683: Is a shift of favour from NHEJ to HR when the cell cycle is progressing from G1 to S/G2 phases in eukaryotic cells. In diploid eukaryotic organisms , the events of meiosis can be viewed as occurring in three steps. (1) Haploid gametes undergo syngamy/ fertilisation with the result that chromosome sets of different parental origin come together to share the same nucleus . (2) Homologous chromosomes originating from different cells (i.e. non-sister chromosomes) align in pairs and undergo recombination involving double-strand break repair. (3) Two successive cell divisions (without duplication of chromosomes) result in haploid gametes that can then repeat

2871-437: 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 . Viruses are found in almost every ecosystem on Earth and are the most numerous type of biological entity. Since Dmitri Ivanovsky 's 1892 article describing a non-bacterial pathogen infecting tobacco plants and

2970-400: Is associated with proteins within a central disc structure known as a nucleoid . The nucleoid is surrounded by a membrane and two lateral bodies of unknown function. The virus has an outer envelope with a thick layer of protein studded over its surface. The whole virion is slightly pleomorphic , ranging from ovoid to brick-shaped. Mimivirus is one of the largest characterised viruses, with

3069-439: Is caused by cessation of its normal activities because of suppression by virus-specific proteins, not all of which are components of the virus particle. The distinction between cytopathic and harmless is gradual. Some viruses, such as Epstein–Barr virus , can cause cells to proliferate without causing malignancy, while others, such as papillomaviruses , are established causes of cancer. Some viruses cause no apparent changes to

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3168-553: Is controversy over whether the bornavirus , previously thought to cause neurological diseases in horses, could be responsible for psychiatric illnesses in humans. Double-strand break repair model DSB can occur naturally due to the presence of reactive species generated by metabolism, and various external factors (e.g. ionizing radiation or chemotherapeutic drugs). In mammalian cells, there are numerous cellular processes that induce DSB. Firstly, DNA topological strain from topoisomerase during normal cell growth can cause

3267-512: Is correct. It seems unlikely that all currently known viruses have a common ancestor, and viruses have probably arisen numerous times in the past by one or more mechanisms. The first evidence of the existence of viruses came from experiments with filters that had pores small enough to retain bacteria. In 1892, Dmitri Ivanovsky used one of these filters to show that sap from a diseased tobacco plant remained infectious to healthy tobacco plants despite having been filtered. Martinus Beijerinck called

3366-454: Is due to the higher eukaryote's larger genome size, as it means that more NHEJ related proteins are encoded for NHEJ repair pathways; and a larger genome implies a challenging obstacle to find a homologous template for HR. HR and NHEJ pathways are favoured in various phases of cell cycles for a multitude of factors. As S and G2 phases of the cell cycle generate more chromatids , the increased availability of template access for HR results in

3465-751: Is evidence that ranavirus infections target macrophages as a mechanism for gaining entry to cells. Andrias davidianus ranavirus, isolated from the Chinese giant salamander, encodes a protein (Rad2 homolog) that has a key role in the repair of DNA by homologous recombination and in DNA double-strand break repair . Transmission of ranaviruses is thought to occur by multiple routes, including contaminated soil, direct contact, waterborne exposure, and ingestion of infected tissues during predation , necrophagy or cannibalism . Ranaviruses are relatively stable in aquatic environments, persisting several weeks or longer outside

3564-507: Is first recorded in 1728, long before the discovery of viruses by Dmitri Ivanovsky in 1892. The English plural is viruses (sometimes also vira ), whereas the Latin word is a mass noun , which has no classically attested plural ( vīra is used in Neo-Latin ). The adjective viral dates to 1948. The term virion (plural virions ), which dates from 1959, is also used to refer to

3663-607: Is identical in sequence to the viral mRNA and is thus a coding strand, while negative-sense viral ssDNA is complementary to the viral mRNA and is thus a template strand. Several types of ssDNA and ssRNA viruses have genomes that are ambisense in that transcription can occur off both strands in a double-stranded replicative intermediate. Examples include geminiviruses , which are ssDNA plant viruses and arenaviruses , which are ssRNA viruses of animals. Genome size varies greatly between species. The smallest—the ssDNA circoviruses, family Circoviridae —code for only two proteins and have

3762-430: Is likely that sympatric fish and reptile species may serve as reservoirs for virus as well. Interclass transmission has been proven through the use of mesocosm studies. Gross lesions associated with Ranavirus infection include erythema, generalized swelling, hemorrhage, limb swelling, and swollen and friable livers. [REDACTED] Data related to List of viruses at Wikispecies Viruses A virus

3861-431: Is limited to a single strain of bacteria and they can be used to trace the source of outbreaks of infections by a method called phage typing . The complete set of viruses in an organism or habitat is called the virome ; for example, all human viruses constitute the human virome . A novel virus is one that has not previously been recorded. It can be a virus that is isolated from its natural reservoir or isolated as

3960-454: Is not understood well, however in realistic environmental conditions the T90 value of an FV3-like virus is 1 day. The duration of persistence is likely affected by temperature and microbial conditions. It is unlikely that ranaviruses persist in the environment outside of host species between outbreak events. Researchers have explored several pathogen reservoirs for the virus which might explain how

4059-495: Is reorganized prior to the development of adult tissues. During this time period, amphibians are stressed, and their immune systems are down regulated. This decrease in immune function and warmer environmental temperatures allows for greater viral replication and cellular damage to occur. Across 64 mortality events in the United States 54% were found to occur between June-August. The environmental persistence of Ranaviruses

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4158-405: Is responsible for RAD51 recruitment or stabilization at damage sites. RAD51 plays a major role in homologous recombinational repair of DNA during double strand break repair. In this process, an ATP dependent DNA strand exchange takes place in which a template strand invades base-paired strands of homologous DNA molecules. RAD51 is involved in the search for homology and strand pairing stages of

4257-500: Is said that the accumulation of unrepaired DNA damage may lead to various diseases, including various metabolic syndromes and types of cancers . Some examples of diseases caused by defects of DSB repair mechanisms are listed below: Women tend to live longer than men and the gender gap in life expectancy suggests differences in the ageing process between the sexes. Sex specific differences in DNA double-strand break repair of cycling human lymphocytes during aging were studied. It

4356-502: Is strictly regulated. HR repairs DSB by copying intact and homologous DNA molecules. The blunt ends of the DSB are processed into ssDNA with 3’ extensions, which allows RAD51 recombinase (eukaryotic homologue of prokaryotic RecA ) to bind to it to form a nucleoprotein filament. The function of the filament is to locate the template DNA and form a joint heteroduplex molecule. Other proteins such as RP-A protein and RAD52 also coordinate in

4455-456: Is the most preferred repair mechanism in somatic cells . The pathway of SDSA is similar to DSBR until just after the D-loop formation. Instead of forming Holliday junctions after DNA synthesis, the nascent strand dissociates via RETL1 helicase and anneals back to the other end of the resected strand. This explains why SDSA results in a non-crossover pathway. The remaining gap is filled in and

4554-475: Is through disease-bearing organisms known as vectors : for example, viruses are often transmitted from plant to plant by insects that feed on plant sap , such as aphids ; and viruses in animals can be carried by blood-sucking insects. Many viruses spread in the air by coughing and sneezing, including influenza viruses , SARS-CoV-2 , chickenpox , smallpox , and measles . Norovirus and rotavirus , common causes of viral gastroenteritis , are transmitted by

4653-477: The CD4 molecule—a chemokine receptor —which is most commonly found on the surface of CD4+ T-Cells . This mechanism has evolved to favour those viruses that infect only cells in which they are capable of replication. Attachment to the receptor can induce the viral envelope protein to undergo changes that result in the fusion of viral and cellular membranes, or changes of non-enveloped virus surface proteins that allow

4752-472: The ERCC1 -XPF complex activity. It is only after the removal of 3’ ssDNA, where the polymerase will fill the missing gaps and the ligase to ligate the strands. Since SSA results in the deletion of repetitive sequences, this could potentially lead to error-prone repair. Single-strand annealing differs from SDSA and DSBR in numerous ways. For instance, the 3’ extension after the end resection in SSA anneals to

4851-555: The International Committee on Taxonomy of Viruses (ICTV) was formed. The system proposed by Lwoff, Horne and Tournier was initially not accepted by the ICTV because the small genome size of viruses and their high rate of mutation made it difficult to determine their ancestry beyond order. As such, the Baltimore classification system has come to be used to supplement the more traditional hierarchy. Starting in 2018,

4950-639: The Latin vīrus , which refers to poison and other noxious liquids. Vīrus comes from the same Indo-European root as Sanskrit viṣa , Avestan vīša , and Ancient Greek ἰός ( iós ), which all mean "poison". The first attested use of "virus" in English appeared in 1398 in John Trevisa 's translation of Bartholomeus Anglicus 's De Proprietatibus Rerum . Virulent , from Latin virulentus ('poisonous'), dates to c.  1400 . A meaning of 'agent that causes infectious disease'

5049-490: The NCBI Virus genome database has more than 193,000 complete genome sequences, but there are doubtlessly many more to be discovered. A virus has either a DNA or an RNA genome and is called a DNA virus or an RNA virus , respectively. Most viruses have RNA genomes. Plant viruses tend to have single-stranded RNA genomes and bacteriophages tend to have double-stranded DNA genomes. Viral genomes are circular, as in

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5148-451: The cell nucleus , where viral DNA replication begins via a virally encoded DNA polymerase . Viral DNA then abandons the cell nucleus and begins the second stage of DNA replication in the cytoplasm, ultimately forming DNA concatemers . The viral DNA is then packaged via a headful mechanism into infectious virions. The ranavirus genome, like other iridoviral genomes is circularly permuted and exhibits terminally redundant DNA . There

5247-539: The common cold , influenza , chickenpox , and cold sores . Many serious diseases such as rabies , Ebola virus disease , AIDS (HIV) , avian influenza , and SARS are caused by viruses. The relative ability of viruses to cause disease is described in terms of virulence . Other diseases are under investigation to discover if they have a virus as the causative agent, such as the possible connection between human herpesvirus 6 (HHV6) and neurological diseases such as multiple sclerosis and chronic fatigue syndrome . There

5346-422: The faecal–oral route , passed by hand-to-mouth contact or in food or water. The infectious dose of norovirus required to produce infection in humans is fewer than 100 particles. HIV is one of several viruses transmitted through sexual contact and by exposure to infected blood. The variety of host cells that a virus can infect is called its host range : this is narrow for viruses specialized to infect only

5445-430: The germline of the host organisms, by which they can be passed on vertically to the offspring of the host for many generations. This provides an invaluable source of information for paleovirologists to trace back ancient viruses that existed as far back as millions of years ago. There are three main hypotheses that aim to explain the origins of viruses: In the past, there were problems with all of these hypotheses:

5544-408: The polyomaviruses , or linear, as in the adenoviruses . The type of nucleic acid is irrelevant to the shape of the genome. Among RNA viruses and certain DNA viruses, the genome is often divided into separate parts, in which case it is called segmented. For RNA viruses, each segment often codes for only one protein and they are usually found together in one capsid. All segments are not required to be in

5643-447: The three domains . This discovery has led modern virologists to reconsider and re-evaluate these three classical hypotheses. The evidence for an ancestral world of RNA cells and computer analysis of viral and host DNA sequences give a better understanding of the evolutionary relationships between different viruses and may help identify the ancestors of modern viruses. To date, such analyses have not proved which of these hypotheses

5742-432: The 5' to 3' cutting of DNA ends, annealing of microhomology, removing heterologous flaps, and ligation and synthesis of gap filling DNA. It was found that the selection between MMEJ and NHEJ is mainly dependent on Ku levels and the concurrent cell cycle. DNA damage response (DDR) is the overarching mechanism which mediates the cell's detection and response to DNA damage. This includes the process of detecting DSB within

5841-584: The DDR, which plays a vital role in DSB repair pathways regulation. The image in this section illustrates molecular steps in a DNA damage response pathway in which a Fanconi anemia complex is activated during repair of a double-strand break. ATM (ATM) is also a protein kinase that is recruited and activated by DNA double-strand breaks . DNA double-strand damages activate the Fanconi anemia core complex (FANCA/B/C/E/F/G/L/M). The FA core complex monoubiquitinates

5940-470: The DSB site in the DNA. In particular, ATM has been identified as the protein kinase in charge of the global meditation of cellular responses to DSB, which includes various DSB repair pathways. Following the recruitment of the aforementioned proteins to DNA damage sites, they will in turn trigger cellular responses and repair pathways to mitigate and repair the damage caused. In short, these vital upstream proteins and downstream repair pathways altogether forms

6039-527: The ICTV began to acknowledge deeper evolutionary relationships between viruses that have been discovered over time and adopted a 15-rank classification system ranging from realm to species. Additionally, some species within the same genus are grouped into a genogroup . The ICTV developed the current classification system and wrote guidelines that put a greater weight on certain virus properties to maintain family uniformity. A unified taxonomy (a universal system for classifying viruses) has been established. Only

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6138-558: The Megalocytiviruses, are an emerging group of closely related dsDNA viruses which cause systemic infections in a wide variety of wild and cultured fresh and saltwater fishes. As with Megalocytiviruses, Ranavirus outbreaks are therefore of considerable economic importance in aquaculture , as epizootics can result in moderate fish loss or mass mortality events of cultured fishes. Unlike Megalocytiviruses, however, Ranavirus infections in amphibians have been implicated as

6237-489: The basis of similarities. In 1962, André Lwoff , Robert Horne , and Paul Tournier were the first to develop a means of virus classification, based on the Linnaean hierarchical system. This system based classification on phylum , class , order , family , genus , and species . Viruses were grouped according to their shared properties (not those of their hosts) and the type of nucleic acid forming their genomes. In 1966,

6336-448: The cell, and the subsequent triggering and regulation of DSB repair pathways. Upstream detections of DNA damage via DDR will lead to the activation of downstream responses such as senescence , cell apoptosis , halting transcription and activating DNA repair mechanisms. Proteins such as the proteins ATM , ATR and DNA-dependent protein kinase (DNA-PK) are vital for the process of detection of DSB in DDR, and these proteins are recruited to

6435-481: The cleavage determines whether the resolution results in either cross-over or noncross-over products. Lastly, the strands finally separate and revert to its original form. , the main pathway for resolution relies on the BTR (BLM helicase-TopoisomeraseIIIα-RMI1-RM2) complex, where it induces the resolution of the 2 Holliday junctions, but this pathway favors the noncross-over cleavage. Synthesis-dependent strain annealing

6534-410: The contrast between viruses and the background, electron-dense "stains" are used. These are solutions of salts of heavy metals, such as tungsten , that scatter the electrons from regions covered with the stain. When virions are coated with stain (positive staining), fine detail is obscured. Negative staining overcomes this problem by staining the background only. A complete virus particle, known as

6633-418: The discovery of the tobacco mosaic virus by Martinus Beijerinck in 1898, more than 11,000 of the millions of virus species have been described in detail. The study of viruses is known as virology , a subspeciality of microbiology . When infected, a host cell is often forced to rapidly produce thousands of copies of the original virus. When not inside an infected cell or in the process of infecting

6732-481: The downstream targets FANCD2 and FANCI. ATM activates (phosphorylates) CHEK2 and FANCD2 CHEK2 phosphorylates BRCA1. Ubiquinated FANCD2 complexes with BRCA1 and RAD51 . The PALB2 protein acts as a hub, bringing together BRCA1, BRCA2 and RAD51 at the site of a DNA double-strand break, and also binds to RAD51C, a member of the RAD51 paralog complex RAD51B - RAD51C - RAD51D - XRCC2 (BCDX2). The BCDX2 complex

6831-631: The ends of a DSB is captured by a group of enzymes. The enzymes then form a bridge which connects the DSB ends together, and is lastly followed by religation of the DNA strands. To initiate whole process, the Ku70 / 80 protein complex binds to the damaged ends of the DSB strands. This forms a preliminary scaffold which allows the recruitment of various NHEJ factors, such as the DNA-dependent protein kinase catalytic subunit (DNA-PKcs), DNA Ligase IV and X-ray cross complementing protein 4 (XRCC4) to form

6930-431: The extreme of the ssRNA virus case. Viruses undergo genetic change by several mechanisms. These include a process called antigenic drift where individual bases in the DNA or RNA mutate to other bases. Most of these point mutations are "silent"—they do not change the protein that the gene encodes—but others can confer evolutionary advantages such as resistance to antiviral drugs . Antigenic shift occurs when there

7029-446: The filtered, infectious substance a "virus" and this discovery is considered to be the beginning of virology. The subsequent discovery and partial characterization of bacteriophages by Frederick Twort and Félix d'Herelle further catalyzed the field, and by the early 20th century many viruses had been discovered. In 1926, Thomas Milton Rivers defined viruses as obligate parasites. Viruses were demonstrated to be particles, rather than

7128-741: The first isolation of a Ranavirus in 1960s from the Northern leopard frog ( Lithobates pipiens ). The ranaviruses appear to have evolved from a fish virus which subsequently infected amphibians and reptiles. The genus contains the following species: The family Iridoviridae is divided into seven genera which include Chloriridovirus , Iridovirus , Lymphocystivirus , Megalocytivirus , and Ranavirus . The genus Ranavirus contains three viruses known to infect amphibians ( Ambystoma tigrinum virus (ATV), Bohle iridovirus (BIV), and frog virus 3 ). Ranaviruses are large icosahedral DNA viruses measuring approximately 150 nm in diameter with

7227-399: The form of single-stranded nucleoprotein complexes, through pores called plasmodesmata . Bacteria, like plants, have strong cell walls that a virus must breach to infect the cell. Given that bacterial cell walls are much thinner than plant cell walls due to their much smaller size, some viruses have evolved mechanisms that inject their genome into the bacterial cell across the cell wall, while

7326-623: The heteroduplex formation, the RP-A protein has to be removed for the RAD51 to form the filament, whereas the RAD52 is a key HR mediator. Afterwards, the 3’ ssDNA invades the template DNA, and displaces a DNA strand to form a D-loop. DNA polymerase and other accessory factors follows by replacing the missing DNA via DNA synthesis. Ligase then attaches the DNA strand break, resulting in the formation of 2 Holliday junctions . The recombined DNA strands then undergoes resolution by cleavage. The orientation of

7425-406: The host. At some point, the provirus or prophage may give rise to the active virus, which may lyse the host cells. Enveloped viruses (e.g., HIV) typically are released from the host cell by budding . During this process, the virus acquires its envelope, which is a modified piece of the host's plasma or other, internal membrane. The genetic material within virus particles, and the method by which

7524-517: The infected cell. Cells in which the virus is latent and inactive show few signs of infection and often function normally. This causes persistent infections and the virus is often dormant for many months or years. This is often the case with herpes viruses . Viruses are by far the most abundant biological entities on Earth and they outnumber all the others put together. They infect all types of cellular life including animals, plants, bacteria and fungi . Different types of viruses can infect only

7623-883: The majority a cell’s DSB. Secondly, cellular processes such as meiosis and the maturation of antibodies can cause nuclease-induced DSB. Thirdly, the cleavage of different DNA structures such as reversed or blocked DNA replication forks , R-loops and DNA interstrand crosslinks can also cause DSB. Homologous recombination involves the exchange of DNA materials between homologous chromosomes. There are multiple pathways of HR to repair DSBs, which includes double-strand break repair (DSBR), synthesis-dependent strand annealing (SDSA), break-induced replication (BIR), and single-strand annealing (SSA). The regulation of HR in mammalian cells involves key HR proteins such as BRCA1 and BRCA2 . And as mentioned, since HR can lead to aggressive chromosomal rearrangement, loss of genetic information that could contribute to cell death, it explains why HR

7722-416: The material is replicated, varies considerably between different types of viruses. The range of structural and biochemical effects that viruses have on the host cell is extensive. These are called ' cytopathic effects '. Most virus infections eventually result in the death of the host cell. The causes of death include cell lysis, alterations to the cell's surface membrane and apoptosis . Often cell death

7821-412: The meiotic cycle. During step (2), damages in DNA of the germline can be removed by double-strand break repair. In particular, double-strand breaks in one duplex DNA molecule can be accurately repaired using information from a homologous intact DNA molecule by the process of homologous recombination . Although there is no universal model to explain disease etiology caused by DNA repair deficiency, it

7920-439: The nick is attached by the ligase. Although there is little research in regards of break-induced replication, it is known that it is a one-ended recombination mechanism, where only of the one ends of a DSB will be involved in strand invasion. This means that unlike DSBR, BIR does not link back to the second DSB end after the strand invasion and replication. Single-strand annealing involves homologous/repeated sequences flanking

8019-461: The original virus. Their life cycle differs greatly between species, but there are six basic stages in their life cycle: Attachment is a specific binding between viral capsid proteins and specific receptors on the host cellular surface. This specificity determines the host range and type of host cell of a virus. For example, HIV infects a limited range of human leucocytes . This is because its surface protein, gp120 , specifically interacts with

8118-432: The other hand, DSB due to replication fork collapse mainly favours HR. It is said that the favoured pathway in a particular situations is also largely dependent on the species of the cell, the cell type, and cell cycle phases; and are all modulated and triggered by different upstream regulatory proteins. As compared to higher eukaryotes , yeast cells have adopted HR as the main repair pathway for DSB. Imprecise NHEJ,

8217-416: The polymerase during genome replication. This process appears to be an adaptation for coping with genome damage. Viral populations do not grow through cell division, because they are acellular. Instead, they use the machinery and metabolism of a host cell to produce multiple copies of themselves, and they assemble in the cell. When infected, the host cell is forced to rapidly produce thousands of copies of

8316-432: The primary pathway for NHEJ to repair "dirty" ends due to IR, was found to be inefficient at repairing DSB in yeast cells. It was hypothesized that this inefficiency as compared to mammalian cells is due to the lack of three vital NHEJ proteins, including DNA-PKcs , BRCA1 , and Artemis . Contrary to yests, higher eukaryotes has a much higher frequency and efficiency at adopting NHEJ pathways. Research hypothesize that this

8415-604: The process. As cells have developed various DSB repair models, it is said that specific pathways are favoured for their ability to repair DSB depending on the cellular context. These conditions include the type of DSB involved, the species of cells involved, and the stage of the cell cycle. Cells have evolved a multitude of DSB repair pathways in response to the various types of DSB. Hence, various pathways are favoured in different situations. For instance, frank DSB, which are DSB induced by substances like as ionizing radiation , and nucleases , can be repaired by both HR and NHEJ. On

8514-965: The ranks of subrealm, subkingdom, and subclass are unused, whereas all other ranks are in use. The Nobel Prize-winning biologist David Baltimore devised the Baltimore classification system. The ICTV classification system is used in conjunction with the Baltimore classification system in modern virus classification. The Baltimore classification of viruses is based on the mechanism of mRNA production. Viruses must generate mRNAs from their genomes to produce proteins and replicate themselves, but different mechanisms are used to achieve this in each virus family. Viral genomes may be single-stranded (ss) or double-stranded (ds), RNA or DNA, and may or may not use reverse transcriptase (RT). In addition, ssRNA viruses may be either sense (+) or antisense (−). This classification places viruses into seven groups: Examples of common human diseases caused by viruses include

8613-422: The regressive hypothesis did not explain why even the smallest of cellular parasites do not resemble viruses in any way. The escape hypothesis did not explain the complex capsids and other structures on virus particles. The virus-first hypothesis contravened the definition of viruses in that they require host cells. Viruses are now recognised as ancient and as having origins that pre-date the divergence of life into

8712-419: The repeated/homologous sequences of the other end, whereas in other pathways the strand invasion to another homologous DNA template. Moreover, SSA does not require RAD51 , because it does not involve strand invasion, but rather the annealing of homologous sequences. Non-homologous end joining (NHEJ) is one of the major pathways in DSB repair besides HR. The basic concept of NHEJ involves three steps. First,

8811-484: The result of spread to an animal or human host where the virus had not been identified before. It can be an emergent virus , one that represents a new virus, but it can also be an extant virus that has not been previously identified . The SARS-CoV-2 coronavirus that caused the COVID-19 pandemic is an example of a novel virus. Classification seeks to describe the diversity of viruses by naming and grouping them on

8910-451: The same virion for the virus to be infectious, as demonstrated by brome mosaic virus and several other plant viruses. A viral genome, irrespective of nucleic acid type, is almost always either single-stranded (ss) or double-stranded (ds). Single-stranded genomes consist of an unpaired nucleic acid, analogous to one-half of a ladder split down the middle. Double-stranded genomes consist of two complementary paired nucleic acids, analogous to

9009-462: The size of most bacteria. The origins of viruses in the evolutionary history of life are still unclear. Some viruses may have evolved from plasmids , which are pieces of DNA that can move between cells. Other viruses may have evolved from bacteria. In evolution, viruses are an important means of horizontal gene transfer , which increases genetic diversity in a way analogous to sexual reproduction . Viruses are considered by some biologists to be

9108-399: The structure-mediated self-assembly of the virus particles, some modification of the proteins often occurs. In viruses such as HIV, this modification (sometimes called maturation) occurs after the virus has been released from the host cell. Release – Viruses can be released from the host cell by lysis , a process that kills the cell by bursting its membrane and cell wall if present: this

9207-457: The study of the origin of life , as it lends further credence to the hypothesis that life could have started as self-assembling organic molecules . The virocell model first proposed by Patrick Forterre considers the infected cell to be the "living form" of viruses and that virus particles (virions) are analogous to spores . Although the living versus non-living debate continues, the virocell model has gained some acceptance. Viruses display

9306-586: The up-regulation of the pathway. This rise is further increased due to the activation of CDK1 and the increase of RAD51 and RAD52 levels during G1 phase. Despite this, NHEJ not is inactive during the HR up-regulation. In fact, NHEJ was shown to be active throughout all stages of the cell cycle, and is favoured in G1 phase during low resection action intervals. This suggests the competition between HR and NHEJ for DSB repair in cells. It should be noted, however, that there

9405-446: The viral messenger RNA (mRNA). Positive-sense viral RNA is in the same sense as viral mRNA and thus at least a part of it can be immediately translated by the host cell. Negative-sense viral RNA is complementary to mRNA and thus must be converted to positive-sense RNA by an RNA-dependent RNA polymerase before translation. DNA nomenclature for viruses with genomic ssDNA is similar to RNA nomenclature, in that positive-strand viral ssDNA

9504-824: The viral capsid remains outside. Uncoating is a process in which the viral capsid is removed: This may be by degradation by viral enzymes or host enzymes or by simple dissociation; the end-result is the releasing of the viral genomic nucleic acid. Replication of viruses involves primarily multiplication of the genome. Replication involves the synthesis of viral messenger RNA (mRNA) from "early" genes (with exceptions for positive-sense RNA viruses), viral protein synthesis , possible assembly of viral proteins, then viral genome replication mediated by early or regulatory protein expression. This may be followed, for complex viruses with larger genomes, by one or more further rounds of mRNA synthesis: "late" gene expression is, in general, of structural or virion proteins. Assembly – Following

9603-403: The virus can persist within an amphibian community. In some amphibian populations, sub-clinically infected individuals may serve as reservoirs for the pathogen. These sub-clinically infected individuals are responsible for reintroduction of the virus to the larval population. With ranaviruses being capable of infected multiple taxa, and with there being differences in susceptibility between taxa, it

9702-503: The virus to enter. Penetration or viral entry follows attachment: Virions enter the host cell through receptor-mediated endocytosis or membrane fusion . The infection of plant and fungal cells is different from that of animal cells. Plants have a rigid cell wall made of cellulose , and fungi one of chitin, so most viruses can get inside these cells only after trauma to the cell wall. Nearly all plant viruses (such as tobacco mosaic virus) can also move directly from cell to cell, in

9801-460: The virus useless or uncompetitive. To compensate, RNA viruses often have segmented genomes—the genome is split into smaller molecules—thus reducing the chance that an error in a single-component genome will incapacitate the entire genome. In contrast, DNA viruses generally have larger genomes because of the high fidelity of their replication enzymes. Single-strand DNA viruses are an exception to this rule, as mutation rates for these genomes can approach

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