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Varidnaviria

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88-481: Varidnaviria is a realm of viruses that includes all DNA viruses that encode major capsid proteins that contain a vertical jelly roll fold . The major capsid proteins (MCP) form into pseudohexameric subunits of the viral capsid , which stores the viral deoxyribonucleic acid (DNA), and are perpendicular, or vertical, to the surface of the capsid. Apart from this, viruses in the realm also share many other characteristics, such as minor capsid proteins (mCP) with

176-580: A body plan from a common ancestor, and that taxa were branches of a single tree of life . The word homology, coined in about 1656, is derived from the Greek ὁμόλογος homologos from ὁμός homos 'same' and λόγος logos 'relation'. Similar biological structures or sequences in different taxa are homologous if they are derived from a common ancestor . Homology thus implies divergent evolution . For example, many insects (such as dragonflies ) possess two pairs of flying wings . In beetles ,

264-404: A common ancestor after having been subjected to adaptive modifications for different purposes as the result of natural selection . The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin 's theory of evolution in 1859, but had been observed before this from Aristotle's biology onwards, and it

352-461: A parvovirus , both of which are classified in the realm Monodnaviria . However, this scenario was contradicted by a molecular phylogenetic analysis suggesting that tectiviruses and polintons had no involvement in the origin of eukaryotic viruses in Varidnaviria and that polintons are probably derived from these eukaryotic viruses. While the jelly roll fold is found in other realms, including

440-577: A reverse transcriptase (RT), assigned to the kingdom Pararnavirae . These enzymes are vital in the viral life cycle, as RdRp transcribes viral mRNA and replicates the genome, and RT likewise replicates the genome. Riboviria mostly contains eukaryotic viruses, and most eukaryotic viruses, including most human, animal, and plant viruses, belong to the realm. Most widely known viral diseases are caused by viruses in Riboviria , which includes influenza viruses , HIV , coronaviruses , ebolaviruses , and

528-524: A tectivirus or tectivirus-like virus of a bacterium that became a bacterial symbiont in a proto-eukaryote. From there, based on phylogenetic analysis of the viral DNA polymerase and other characteristics, eukaryotic viruses in Bamfordvirae appear to have formed a complex relationship with various selfish genetic elements , including polintons , a type of transposon , portions of DNA that can self-replicate and integrate themselves into other parts of

616-506: A P-loop fold at the end of the protein's amino acid sequence , and the HerA family is homologous to FtsK. The exact function of the ATPase for some viruses in Varidnaviria is unclear since morphological features, such as the circular, supercoiled genome of Pseudoalteromonas virus PM2 , seemingly prohibit translocation by the ATPase of DNA from outside the capsid to the inside. The subset of

704-404: A Swiss roll. Each beta strand is a specific sequence of amino acids , and these strands bond to their antiparallel strands via hydrogen bonds . The difference between SJR and DJR folds is that a DJR fold is simply two SJR folds in a single protein. Vertical folds are those that are perpendicular to the capsid surface, in contrast to horizontal folds that are parallel to the capsid surface. During

792-426: A capsid that is made of major capsid proteins that contain vertical single (SJR) or double jelly roll (DJR) folds. The major capsid proteins are named so because they are the primary proteins that the capsid is made of. A jelly roll fold is a type of folded structure in a protein in which eight antiparallel beta strands are organized into four antiparallel beta sheets in a layout resembling a jelly roll , also called

880-569: A gene fusion event that merged the two SJR-MCPs into one, indicated by the two SJR-MCPs forming a lattice in the capsid that structurally resembles the DJR-MCP capsid lattice. Archaeal dsDNA viruses in Portogloboviridae contain just one vertical SJR-MCP, which appears to have been duplicated to two for Halopanivirales , so the MCP of Portogloboviridae likely represents an earlier stage in

968-470: A gene in an organism is duplicated, the two copies are paralogous. They can shape the structure of whole genomes and thus explain genome evolution to a large extent. Examples include the Homeobox ( Hox ) genes in animals. These genes not only underwent gene duplications within chromosomes but also whole genome duplications . As a result, Hox genes in most vertebrates are spread across multiple chromosomes:

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1056-541: A highly derived offshoot of the Nucleocytoviricota . Bacteriophages in Varidnaviria are potentially a major cause of death among marine prokaryotes . This viewpoint is based on autolykiviruses having broad host ranges, infecting and killing many different strains of various bacteria species, in contrast to tailed bacteriophages, which have more limited host ranges, as well as on the apparently large number of marine non-tailed dsDNA viruses. Algal viruses of

1144-409: A much larger number of genes than typical viruses. The name " Varidnaviria " is a portmanteau of vari ous DNA viruses and the suffix - viria , which is the suffix used for virus realms. Double-stranded DNA (dsDNA) viruses in the realm are often called non-tailed or tailless dsDNA viruses to distinguish them from the tailed dsDNA viruses of Duplodnaviria . Most viruses in Varidnaviria contain

1232-480: A pattern of gene expression in the growing zones ( meristems ) is described by the ABC model of flower development . Each of the four types of flower parts is serially repeated in concentric whorls, controlled by a small number of genes acting in various combinations. Thus, A genes working alone result in sepal formation; A and B together produce petals; B and C together create stamens; C alone produces carpels. When none of

1320-459: A single vertical jelly roll (SJR) fold in the MCP, assigned to the kingdom Helvetiavirae . The DJR-MCP lineage is thought to be descended from the SJR-MCP lineage via a gene fusion event, and the SJR-MCP shows a close relation to nucleoplasmins , pointing to a possible origin of the realm's jelly roll fold MCP. Most identified eukaryotic DNA viruses belong to Varidnaviria . Marine viruses in

1408-462: A vast undescribed diversity of viruses in this part of the virosphere. Ribozyviria is characterised by the presence of genomic and antigenomic ribozymes of the Deltavirus type. Additional common features include a rod-like structure and a RNA-binding "delta antigen" encoded in the genome. In general, virus realms have no genetic relation to each other based on common descent, in contrast to

1496-487: Is a synapomorphy for fleas. Patterns such as these lead many cladists to consider the concept of homology and the concept of synapomorphy to be equivalent. Some cladists follow the pre-cladistic definition of homology of Haas and Simpson, and view both synapomorphies and symplesiomorphies as homologous character states. Homologies provide the fundamental basis for all biological classification, although some may be highly counter-intuitive. For example, deep homologies like

1584-467: Is composed of asymmetric units containing two MCP molecules, a homodimer in the case of rudivirids and a heterodimer of paralogous MCPs in the case of lipothrixvirids and tristromavirids. The MCPs of ligamenviral particles have a unique α-helical fold first found in the MCP of rudivirid Sulfolobus islandicus rod-shaped virus 2 (SIRV2). All members of the Adnaviria share a characteristic feature in that

1672-549: Is defined in terms of shared ancestry. Two segments of DNA can have shared ancestry because of either a speciation event ( orthologs ) or a duplication event ( paralogs ). Homology among proteins or DNA is typically inferred from their sequence similarity. Significant similarity is strong evidence that two sequences are related by divergent evolution of a common ancestor. Alignments of multiple sequences are used to indicate which regions of each sequence are homologous. Homologous sequences are orthologous if they are descended from

1760-443: Is shared due to common ancestry. Primary homology may be conceptually broken down further: we may consider all of the states of the same character as "homologous" parts of a single, unspecified, transformation series. This has been referred to as topographical correspondence. For example, in an aligned DNA sequence matrix, all of the A, G, C, T or implied gaps at a given nucleotide site are homologous in this way. Character state identity

1848-404: Is strong evidence that two sequences are related by divergent evolution from a common ancestor. Alignments of multiple sequences are used to discover the homologous regions. Homology remains controversial in animal behaviour , but there is suggestive evidence that, for example, dominance hierarchies are homologous across the primates . Homology was noticed by Aristotle (c. 350 BC), and

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1936-473: Is taken from ribo nucleic acid (RNA), and the first part of Varidnaviria means "various DNA". For viroids , the suffix is designated as - viroidia , and for satellites , the suffix is - satellitia , but as of 2019 neither viroid nor satellite realms have been designated. Duplodnaviria contains double-stranded DNA (dsDNA) viruses that encode a major capsid protein (MCP) that has the HK97 fold. Viruses in

2024-450: Is taken to be homologous. As implied in this definition, many cladists consider secondary homology to be synonymous with synapomorphy , a shared derived character or trait state that distinguishes a clade from other organisms. Shared ancestral character states, symplesiomorphies, represent either synapomorphies of a more inclusive group, or complementary states (often absences) that unite no natural group of organisms. For example,

2112-502: Is the highest taxonomic rank established for viruses by the International Committee on Taxonomy of Viruses (ICTV), which oversees virus taxonomy. Six virus realms are recognized and united by specific highly conserved traits: The rank of realm corresponds to the rank of domain used for cellular life, but differs in that viruses in a realm do not necessarily share a common ancestor based on common descent nor do

2200-404: Is the hypothesis that the particular condition in two or more taxa is "the same" as far as our character coding scheme is concerned. Thus, two Adenines at the same aligned nucleotide site are hypothesized to be homologous unless that hypothesis is subsequently contradicted by other evidence. Secondary homology is implied by parsimony analysis , where a character state that arises only once on a tree

2288-461: Is used as the vaccine against smallpox. ASFV is usually asymptomatic in its natural reservoirs but causes a lethal hemorrhagic fever in domestic pigs that is a concern for agricultural production. Many viruses in Varidnaviria encode the enzyme integrase, allowing them to integrate their genome into their host and behave like transposons. The closely related polintons are apparently always endogenized in their hosts. This integration of viral DNA into

2376-490: The Nucleocytoviricota and thus could be highly derived members of the DJR-MCP viruses, despite the absence of the DJR-MCP and formation of odd-shaped virions. Preliminary phylogenetic analysis of several essential genes that are shared by all these arthropod viruses and the Nucleocytoviricota , such as PolB, RNAP subunits, helicase-primase and thiol oxidoreductase, has suggested that this group of viruses might be

2464-594: The pax6 genes that control the development of the eyes of vertebrates and arthropods were unexpected, as the organs are anatomically dissimilar and appeared to have evolved entirely independently. The embryonic body segments ( somites ) of different arthropod taxa have diverged from a simple body plan with many similar appendages which are serially homologous, into a variety of body plans with fewer segments equipped with specialised appendages. The homologies between these have been discovered by comparing genes in evolutionary developmental biology . Among insects,

2552-532: The African swine fever virus (ASFV). Adenoviruses typically cause mild respiratory, gastrointestinal, and conjunctival illnesses, but occasionally cause more severe illnesses, such as hemorrhagic cystitis , hepatitis , and meningoencephalitis . Poxviruses infect many animals and typically cause non-specific symptoms paired with a characteristic rash that is called a pox. Notable poxviruses include Variola virus , which causes smallpox, and Vaccinia virus , which

2640-412: The embryo in the same manner and from similar origins, such as from matching primordia in successive segments of the same animal, are serially homologous . Examples include the legs of a centipede , the maxillary and labial palps of an insect , and the spinous processes of successive vertebrae in a vertebrate's backbone . Male and female reproductive organs are homologous if they develop from

2728-413: The rabies virus , as well as the first virus to be discovered, Tobacco mosaic virus . Reverse transcribing viruses are a major source of horizontal gene transfer by means of becoming endogenized in their host's genome, and a significant portion of the human genome consists of this viral DNA. Varidnaviria contains DNA viruses that encode MCPs that have a jelly roll fold folded structure in which

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2816-471: The stinger of the female honey bee is a modified ovipositor , homologous with ovipositors in other insects such as the Orthoptera , Hemiptera , and those Hymenoptera without stingers. The three small bones in the middle ear of mammals including humans, the malleus , incus , and stapes , are today used to transmit sound from the eardrum to the inner ear . The malleus and incus develop in

2904-423: The "same organ in different animals under every variety of form and function", and contrasting it with the matching term "analogy" which he used to describe different structures with the same function. Owen codified 3 main criteria for determining if features were homologous: position, development, and composition. In 1859, Charles Darwin explained homologous structures as meaning that the organisms concerned shared

2992-648: The FtsK-HerA superfamily found in Varidnaviria is often called the A32 clade, named after the ATPase-encoding A32(R) gene of Vaccinia virus . Apart from the core morphogenetic triad of genes, the MCP, mCP, and ATPase, certain other characteristics are common or unique in various lineages within Varidnaviria , listed hereafter. It has been suggested that Varidnaviria predates the last universal common ancestor (LUCA) of cellular life and that viruses in

3080-496: The HoxA–D clusters are the best studied. Some sequences are homologous, but they have diverged so much that their sequence similarity is not sufficient to establish homology. However, many proteins have retained very similar structures, and structural alignment can be used to demonstrate their homology. It has been suggested that some behaviours might be homologous, based either on sharing across related taxa or on common origins of

3168-462: The atypical members of Monodnaviria . Eukaryotic monodnaviruses are associated with many diseases, and they include papillomaviruses and polyomaviruses , which cause many cancers, and geminiviruses , which infect many economically important crops. Riboviria contains all RNA viruses that encode an RNA-dependent RNA polymerase (RdRp), assigned to the kingdom Orthornavirae , and all reverse transcribing viruses, i.e. all viruses that encode

3256-538: The behaviour in an individual's development; however, the notion of homologous behavior remains controversial, largely because behavior is more prone to multiple realizability than other biological traits. For example, D. W. Rajecki and Randall C. Flanery, using data on humans and on nonhuman primates , argue that patterns of behaviour in dominance hierarchies are homologous across the primates. As with morphological features or DNA, shared similarity in behavior provides evidence for common ancestry. The hypothesis that

3344-580: The desire to establish higher-level taxonomy for viruses. In two votes in 2018 and 2019, the ICTV agreed to adopt a 15-rank classification system for viruses, ranging from realm to species. Riboviria was established in 2018 based on phylogenetic analysis of the RNA-dependent polymerases being monophyletic, Duplodnaviria was established in 2019 based on increasing evidence that tailed bacteriophages and herpesviruses shared many traits, Monodnaviria

3432-482: The embryo from structures that form jaw bones (the quadrate and the articular) in lizards, and in fossils of lizard-like ancestors of mammals. Both lines of evidence show that these bones are homologous, sharing a common ancestor. Among the many homologies in mammal reproductive systems , ovaries and testicles are homologous. Rudimentary organs such as the human tailbone , now much reduced from their functional state, are readily understood as signs of evolution ,

3520-516: The embryos develop. The implication that the ancestors of snakes had hind legs is confirmed by fossil evidence: the Cretaceous snake Pachyrhachis problematicus had hind legs complete with hip bones ( ilium , pubis , ischium ), thigh bone ( femur ), leg bones ( tibia , fibula ) and foot bones ( calcaneum , astragalus ) as in tetrapods with legs today. As with anatomical structures, sequence homology between protein or DNA sequences

3608-401: The establishment of Riboviria in 2018, three realms in 2019, and two in 2020. The names of realms consist of a descriptive first part and the suffix - viria , which is the suffix used for virus realms. The first part of Duplodnaviria means "double DNA", referring to dsDNA viruses, the first part of Monodnaviria means "single DNA", referring to ssDNA viruses, the first part of Riboviria

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3696-506: The evolutionary history of Varidnaviria MCPs. However, another scenario was later proposed in which the Bamfordvirae and Helvetiavirae kingdoms would originate independently suggesting that the Bamfordvirae DJR-MCP protein snow a relation with the bacterial DUF 2961 protein, leading to a revision of the realm Varidnaviria . It is possible that the Bamfordvirae DJR-MCP will evolve from this protein independently, however

3784-472: The explanation being that they were cut down by natural selection from functioning organs when their functions were no longer needed, but make no sense at all if species are considered to be fixed. The tailbone is homologous to the tails of other primates. In many plants, defensive or storage structures are made by modifications of the development of primary leaves , stems , and roots . Leaves are variously modified from photosynthetic structures to form

3872-546: The family Microviridae in Monodnaviria and various single-stranded RNA viruses in Riboviria , the jelly roll fold found in Varidnaviria is vertical, i.e. perpendicular to the capsid surface, contrary to the jelly roll folds in other realms, which are horizontal, i.e. parallel to the capsid surface. In general, the other virus realms have no apparent relation based on common descent to Varidnaviria . Varidnaviria has two kingdoms: Bamfordvirae and Helvetiavirae ,

3960-446: The family Phycodnaviridae play an important role in controlling algal blooms as well as, with many marine viruses in general, contributing to a process called viral shunt , whereby organic material from killed organisms are "shunted" by viruses away from higher trophic levels and recycled for consumption by those at lower trophic levels. The most notable disease-causing viruses in Varidnaviria are adenoviruses, poxviruses, and

4048-442: The first SJR-MCP virus discovered. Over time, the use of metagenomics has allowed for the identification of viruses in the environment even without identification of the host or laboratory specimens, leading to the discovery of many additional members of the realm. Morphological surveys of marine samples suggest that non-tailed dsDNA viruses may be more numerous than the tailed dsDNA viruses of Duplodnaviria , which, as of 2019, are

4136-448: The first disease to be eradicated. Human adenoviruses were the first DJR-MCP viruses in Varidnaviria to have their MCPs analyzed, standing out for having jelly roll folds that were perpendicular, rather than parallel, to the capsid surface. In 1999, the structure of the MCP of Pseudomonas virus PRD1 was resolved, showing that the DJR-MCP lineage included prokaryotic viruses. Haloarcula hispanica virus SH1 would later, in 2003, become

4224-460: The first eukaryotic viruses in Bamfordvirae or related to the first ones. Polintons then gave rise to multiple lineages by various mechanisms. Among these lineages are full-fledged viruses, including adenoviruses and giant viruses, cytoplasmic linear plasmids, virophages , which are satellite viruses of giant viruses, transpovirons , which are linear plasmid-like DNA molecules found in giant viruses, and bidnaviruses via genetic recombination with

4312-460: The first pair of wings has evolved into a pair of hard wing covers , while in Dipteran flies the second pair of wings has evolved into small halteres used for balance. Similarly, the forelimbs of ancestral vertebrates have evolved into the front flippers of whales , the wings of birds , the running forelegs of dogs , deer , and horses , the short forelegs of frogs and lizards , and

4400-445: The genes are active, leaves are formed. Two more groups of genes, D to form ovules and E for the floral whorls, complete the model. The genes are evidently ancient, as old as the flowering plants themselves. Developmental biology can identify homologous structures that arose from the same tissue in embryogenesis . For example, adult snakes have no legs, but their early embryos have limb-buds for hind legs, which are soon lost as

4488-561: The grasping hands of primates including humans. The same major forearm bones ( humerus , radius , and ulna ) are found in fossils of lobe-finned fish such as Eusthenopteron . The opposite of homologous organs are analogous organs which do similar jobs in two taxa that were not present in their most recent common ancestor but rather evolved separately . For example, the wings of insects and birds evolved independently in widely separated groups , and converged functionally to support powered flight , so they are analogous. Similarly,

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4576-410: The host's genome is a form of horizontal gene transfer between unrelated organisms, although polintons are typically transmitted vertically from parent to child. A peculiar example of endogenization in Varidnaviria are virophages, satellite viruses that are dependent on giant virus infection to replicate. Virophages replicate by hijacking the replication apparati of giant viruses, thereby suppressing

4664-601: The insect-trapping pitchers of pitcher plants , the insect-trapping jaws of the Venus flytrap , and the spines of cactuses , all homologous. Certain compound leaves of flowering plants are partially homologous both to leaves and shoots, because their development has evolved from a genetic mosaic of leaf and shoot development. The four types of flower parts, namely carpels , stamens , petals , and sepals , are homologous with and derived from leaves, as Goethe correctly noted in 1790. The development of these parts through

4752-456: The interaction between the MCP dimer and the linear dsDNA genome maintains the DNA in the A form. Consequently, the entire genome adopts the A form in virions. Like many structurally related viruses in the two other realms of dsDNA viruses ( Duplodnaviria and Varidnaviria ), there is no detectable sequence similarity among the capsid proteins of viruses from different tokiviricete families, suggesting

4840-651: The jelly roll (JR) fold is perpendicular to the surface of the viral capsid. Many members also share a variety of other characteristics, including a minor capsid protein that has a single JR fold, an ATPase that packages the genome during capsid assembly, and a common DNA polymerase . Two kingdoms are recognized: Helvetiavirae , whose members have MCPs with a single vertical JR fold, and Bamfordvirae , whose members have MCPs with two vertical JR folds. Marine viruses in Varidnaviria are ubiquitous worldwide and, like tailed bacteriophages, play an important role in marine ecology. Most identified eukaryotic DNA viruses belong to

4928-422: The largest and most diverse lineage of viruses documented. With the increased knowledge of the viruses of the realm, Varidnaviria was established in 2019 based on the shared traits of viruses in the realm. The establishment of Varidnaviria has allowed for newly discovered and related, yet divergent, viruses to be classified in higher taxa. This includes proposed families such as Finnlakeviridae , which would be

5016-464: The latter of which is monotypic down to the rank of family. This taxonomy can be visualized as follows: All recognized members of Varidnaviria belong to Group I: dsDNA viruses of the Baltimore classification system, which groups viruses together based on how they produce messenger RNA. The family Finnlakeviridae , a proposed family of Varidnaviria , belongs to Group II: ssDNA viruses and would be

5104-482: The number of giant virus virions produced, increasing the likelihood of host survival. Some virophages are able to become endogenized, and this endogenization can be considered a form of adaptive immunity for the host against giant virus infection. Diseases caused by poxviruses have been known for much of recorded history. Smallpox in particular has been highly prominent in modern medicine; the first vaccine to be invented targeted smallpox, and smallpox would later become

5192-444: The only family in the realm with a single-stranded DNA genome, Autolykiviridae , which have a broad host range and which may play a major role in the deaths of marine bacteria , and the "Odin" group, which encode a protein that has no known relation to any other proteins in place of the FtsK-HerA superfamily ATPase. In 2020, autolykiviruses were officially classified for the first time. Realm (virology) In virology , realm

5280-473: The only ssDNA virus in the realm. Most identified DNA viruses that infect eukaryotes belong to Varidnaviria , the other major lineages of eukaryotic DNA viruses being the order Herpesvirales , which infect animals, in Duplodnaviria , and the class Papovaviricetes , which infect animals, in Monodnaviria . Realms are the highest level of taxonomy used for viruses in and Varidnaviria is one of four,

5368-502: The order Herpesvirales . The relation between caudoviruses and herpesviruses is not certain, as they may either share a common ancestor or herpesviruses may be a divergent clade from within Caudovirales . A common trait among duplodnaviruses is that they cause latent infections without replication while still being able to replicate in the future. Tailed bacteriophages are ubiquitous worldwide, important in marine ecology, and

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5456-495: The origin of the DJR-MCP by duplication of the Helvetiavirae SJR-MCP cannot yet be ruled out. A molecular phylogenetic analysis suggests that Helvetiavirae had no involvement in the origin of the Bamfordvirae DJR-MCP and that they probably derive from the class Tectiliviricetes . Viruses in Bamfordvirae appear to have made crossed from prokaryotes to eukaryotes early in eukaryotic history, via infection by

5544-479: The other three being Duplodnaviria , Monodnaviria , and Riboviria . The unassigned family Portogloboviridae is a proposed family of the realm since its capsid proteins appear to be homologous to those of viruses in Varidnaviria . Another proposed group is the Naldaviricetes class (including Polydnaviridae ). These viruses encompass several genes that are distantly related to core genes of

5632-446: The pentagonal vertices of the icosahedral capsid. Most members of the realm also encode genome packaging ATPases of the FtsK-HerA superfamily. The ATPases in Varidnaviria are enzymes that package the viral DNA into the capsid during the process of assembling virions. FtsK is a family of proteins that contains a transmembrane protein with four membrane-spanning helices at the start of the protein's amino acid sequence and an ATPase with

5720-400: The presence of wings is a synapomorphy for pterygote insects, but a symplesiomorphy for holometabolous insects. Absence of wings in non-pterygote insects and other organisms is a complementary symplesiomorphy that unites no group (for example, absence of wings provides no evidence of common ancestry of silverfish, spiders and annelid worms). On the other hand, absence (or secondary loss) of wings

5808-435: The principle of connections, namely that what is important is the relative position of different structures and their connections to each other. Embryologist Karl Ernst von Baer stated what are now called von Baer's laws in 1828, noting that related animals begin their development as similar embryos and then diverge: thus, animals in the same family are more closely related and diverge later than animals which are only in

5896-467: The process of assembling the viral capsid, MCPs self-assemble into hexagonal structures, called hexons, containing multiple copies of the MCP. Hexons then bond to form the relatively flat triangular sides of the icosahedral capsid. All viruses in Varidnaviria that encode a DJR-MCP that have been analyzed in high resolution also encode a minor capsid protein (mCP) that contains an SJR fold. These mCPs assemble into pentagonal structures called pentons that form

5984-612: The rank of subrealm. Prior to the 21st century, it was believed that deep evolutionary relations between viruses could not be discovered due to their high mutation rates and small number of genes making discovering these relations more difficult. Because of this, the highest taxonomic rank for viruses from 1991 to 2017 was order. In the 21st century, however, various methods have been developed that have enabled these deeper evolutionary relationships to be studied, including metagenomics, which has identified many previously unidentified viruses, and comparison of highly conserved traits, leading to

6072-419: The realm also share a number of other characteristics involving the capsid and capsid assembly, including an icosahedral capsid shape and a terminase enzyme that packages viral DNA into the capsid during assembly. Two groups of viruses are included in the realm: tailed bacteriophages, which infect prokaryotes and are assigned to the order Caudovirales , and herpesviruses, which infect animals and are assigned to

6160-416: The realm are called CRESS-DNA viruses and have circular ssDNA genomes. ssDNA viruses with linear genomes are descended from them, and in turn some dsDNA viruses with circular genomes are descended from linear ssDNA viruses. CRESS-DNA viruses include three kingdoms that infect prokaryotes: Loebvirae , Sangervirae , and Trapavirae . The kingdom Shotokuvirae contains eukaryotic CRESS-DNA viruses and

6248-562: The realm are highly abundant worldwide and are important in marine ecology. Many animal viruses in the realm are associated with disease, including adenoviruses , poxviruses , and the African swine fever virus . Poxviruses have been prominent in the history of medicine, especially smallpox , caused by Variola virus , which was the target of the first vaccine and which later became the first disease eradicated. The realm also notably includes giant viruses that are physically larger and contain

6336-583: The realm were present in the LUCA. The vertical SJR-MCPs of Halopanivirales , assigned to the kingdom Helvetiavirae , unlike SJR folds found outside of Varidnaviria , show a relation to a group of proteins that includes the Cupin superfamily and nucleoplasmins, pointing to a possible origin of the major capsid protein of Varidnaviria among this group. The DJR-MCP lineage, assigned to the kingdom Bamfordvirae , thereafter appears to have come into existence by means of

6424-848: The realm. Notable disease-causing viruses in Varidnaviria include adenoviruses , poxviruses , and the African swine fever virus . Poxviruses have been highly prominent in the history of modern medicine, especially Variola virus , which caused smallpox . Many varidnaviruses are able to become endogenized, and a peculiar example of this are virophages , which confer protection for their hosts against giant viruses during infection. Realm Adnaviria unifies archaeal filamentous viruses with linear A-form double-stranded DNA genomes and characteristic major capsid proteins unrelated to those encoded by other known viruses. The realm currently includes viruses from three families, Lipothrixviridae , Rudiviridae , and Tristromaviridae , all infecting hyperthermophilic archaea. The nucleoprotein helix of adnaviruses

6512-411: The realms generally have no genetic relation to each other, there are some exceptions: In virology, the second highest taxonomy rank established by the ICTV is subrealm, which is the rank below realm. Subrealms of viruses use the suffix - vira , viroid subrealms use the suffix - viroida , and satellites use the suffix - satellitida . The rank below subrealm is kingdom. As of 2019, no taxa are described at

6600-529: The realms share a common ancestor . Instead, realms group viruses together based on specific traits that are highly conserved over time, which may have been obtained on a single occasion or multiple occasions. As such, each realm represents at least one instance of viruses coming into existence. While historically it was difficult to determine deep evolutionary relations between viruses, in the 21st century methods such as metagenomics and cryogenic electron microscopy have enabled such research to occur, which led to

6688-414: The same order and have fewer homologies. Von Baer's theory recognises that each taxon (such as a family) has distinctive shared features, and that embryonic development parallels the taxonomic hierarchy: not the same as recapitulation theory . The term "homology" was first used in biology by the anatomist Richard Owen in 1843 when studying the similarities of vertebrate fins and limbs, defining it as

6776-434: The same DNA molecule, and certain types of plasmids , which are extra-chromosomal DNA molecules that self-replicate inside of the cell or organelle that they occupy. The initial bacterial symbiont is likely to have become mitochondria, with mitochondrial linear plasmids descended from tectiviruses remaining. Another divergent lineage reached the nucleus and recombined with transposons, becoming polintons, which may have been

6864-423: The same ancestral sequence separated by a speciation event: when a species diverges into two separate species, the copies of a single gene in the two resulting species are said to be orthologous . The term "ortholog" was coined in 1970 by the molecular evolutionist Walter Fitch . Homologous sequences are paralogous if they were created by a duplication event within the genome. For gene duplication events, if

6952-444: The same embryonic tissue, as do the ovaries and testicles of mammals , including humans . Sequence homology between protein or DNA sequences is similarly defined in terms of shared ancestry. Two segments of DNA can have shared ancestry because of either a speciation event ( orthologs ) or a duplication event ( paralogs ). Homology among proteins or DNA is inferred from their sequence similarity. Significant similarity

7040-470: The subject of much research. Herpesviruses are known to cause a variety of epithelial diseases, including herpes simplex , chickenpox and shingles , and Kaposi's sarcoma . Monodnaviria contains single-stranded DNA (ssDNA) viruses that encode an endonuclease of the HUH superfamily that initiates rolling circle replication and all other viruses descended from such viruses. The prototypical members of

7128-472: The three domains of cellular life— Archaea , Bacteria , and Eukarya —which share a common ancestor. Likewise, viruses within each realm are not necessarily descended from a common ancestor since realms group viruses together based on highly conserved traits, not common ancestry, which is used as the basis for the taxonomy of cellular life. As such, each virus realm is considered to represent at least one instance of viruses coming into existence. By realm: While

7216-432: The three groups. Thus, in the pterosaurs, the "wing" involves both the forelimb and the hindlimb. Analogy is called homoplasy in cladistics , and convergent or parallel evolution in evolutionary biology. Specialised terms are used in taxonomic research. Primary homology is a researcher's initial hypothesis based on similar structure or anatomical connections, suggesting that a character state in two or more taxa share

7304-429: The vertical jelly roll fold, an ATPase that packages viral DNA into the capsid, and a DNA polymerase that replicates the viral genome. Varidnaviria was established in 2019 based on the shared characteristics of the viruses in the realm. There are two groups of viruses in Varidnaviria : viruses that have a double vertical jelly roll (DJR) folds in the MCP, assigned to the kingdom Bamfordvirae , and viruses that have

7392-436: The wings of a sycamore maple seed and the wings of a bird are analogous but not homologous, as they develop from quite different structures. A structure can be homologous at one level, but only analogous at another. Pterosaur , bird and bat wings are analogous as wings, but homologous as forelimbs because the organ served as a forearm (not a wing) in the last common ancestor of tetrapods , and evolved in different ways in

7480-438: Was described late in the 18th century. The French zoologist Etienne Geoffroy Saint-Hilaire showed in 1818 in his theorie d'analogue ("theory of homologues") that structures were shared between fishes, reptiles, birds, and mammals. When Geoffroy went further and sought homologies between Georges Cuvier 's embranchements , such as vertebrates and molluscs, his claims triggered the 1830 Cuvier-Geoffroy debate . Geoffroy stated

7568-482: Was established in 2019 after the relation and origin of CRESS-DNA viruses was resolved, and Varidnaviria was established 2019 based on the shared characteristics of member viruses. Homolog In biology , homology is similarity in anatomical structures or genes between organisms of different taxa due to shared ancestry , regardless of current functional differences. Evolutionary biology explains homologous structures as retained heredity from

7656-426: Was explicitly analysed by Pierre Belon in 1555. A common example of homologous structures is the forelimbs of vertebrates , where the wings of bats and birds , the arms of primates , the front flippers of whales , and the forelegs of four-legged vertebrates like horses and crocodilians are all derived from the same ancestral tetrapod structure. In developmental biology , organs that developed in

7744-690: Was explicitly analysed by Pierre Belon in his 1555 Book of Birds , where he systematically compared the skeletons of birds and humans. The pattern of similarity was interpreted as part of the static great chain of being through the mediaeval and early modern periods: it was not then seen as implying evolutionary change. In the German Naturphilosophie tradition, homology was of special interest as demonstrating unity in nature. In 1790, Goethe stated his foliar theory in his essay "Metamorphosis of Plants", showing that flower parts are derived from leaves. The serial homology of limbs

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