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<title>Cosmid</title>

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<span id="openzim-page-title" class="mw-page-title-main"><span class="mw-page-title-main">Cosmid</span></span>

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<p>A <b>cosmid</b> is a type of hybrid <a href="Plasmid" title="Plasmid">plasmid</a> that contains a <a href="Lambda_phage" title="Lambda phage">Lambda phage</a> <i>cos</i> sequence.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Often used as <a href="Cloning_vector" title="Cloning vector">cloning vectors</a> in <a href="Genetic_engineering" title="Genetic engineering">genetic engineering</a>, cosmids can be used to build <a href="Library_(biology)" title="Library (biology)">genomic libraries</a>. They were first described by Collins and Hohn in 1978.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>

Cosmids can contain 37 to 52 (normally 45) <a href="Base_pair" title="Base pair">kb</a> of DNA, limits based on the normal bacteriophage packaging size. They can <a href="DNA_replication" title="DNA replication">replicate</a> as plasmids if they have a suitable <a href="Origin_of_replication" title="Origin of replication">origin of replication</a> (ori): for example <a href="SV40" title="SV40">SV40</a> ori in mammalian cells, <a href="ColE1" title="ColE1">ColE1</a> ori for double-stranded DNA replication, or <a href="F1_phage" class="mw-redirect" title="F1 phage">f1</a> ori for single-stranded DNA replication in <a href="Prokaryote" title="Prokaryote">prokaryotes</a>. They frequently also contain a gene for selection such as <a href="Antibiotic_resistance" class="mw-redirect" title="Antibiotic resistance">antibiotic resistance</a>, so that the transformed cells can be identified by plating on a medium containing the antibiotic. Those cells which did not take up the cosmid would be unable to grow.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>

</p><p>Unlike plasmids, they can also be packaged in vitro into <a href="Phage" class="mw-redirect" title="Phage">phage</a> <a href="Capsid" title="Capsid">capsids</a>, a step which requires <i>cohesive ends</i>, also known as <i>cos</i> sites also used in cloning with a lambda phage as a vector, however nearly all the lambda genes have been deleted with the exception of the <i>cos</i> sequence. The hybrid cosmid DNA in the capsids can then be transferred into bacterial cells by <a href="Transduction_(genetics)" title="Transduction (genetics)">transduction</a>. Since there is a requirement for in vitro packaging whereby at least 38 kb of DNA is required between the cos sites, the vector without insert DNA will not be packaged (plasmids instability is increased if the novel inserted DNA contains many direct repeats or palindromic (inverted repeats) DNA. This instability can largely be counteracted by using a host bacterium with specific mutations affecting DNA recombination (N.B. Absence of inverted repeats was noted in the first Hohn & Collins publication cited above; see also<sup id="cite_ref-pmid6271486_4-0" class="reference"><a href="#cite_note-pmid6271486-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup>).

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<div class="mw-heading mw-heading2"><h2 id="Cos_sequences"><i>Cos</i> sequences</h2></div>

<p><i>Cos</i> sequences are ~200 base pairs long and essential for packaging. They contain a <i>cosN</i> site where DNA is nicked at each strand, 12 bp apart, by terminase. This causes linearization of the circular cosmid with two "cohesive" or "sticky ends" of 12bp. (The DNA must be linear to fit into a phage head.) The <i>cosB</i> site holds the terminase while it is nicking and separating the strands. The <i>cosQ</i> site of next cosmid (as <a href="Rolling_circle_replication" title="Rolling circle replication">rolling circle replication</a> often results in linear <a href="Concatemers" class="mw-redirect" title="Concatemers">concatemers</a>) is held by the terminase after the previous cosmid has been packaged, to prevent degradation by cellular <a href="DNase" class="mw-redirect" title="DNase">DNases</a>.

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<div class="mw-heading mw-heading2"><h2 id="Cosmid_features_and_uses">Cosmid features and uses</h2></div>

<p>Cosmids are predominantly plasmids with a bacterial <i>oriV</i>, an antibiotic selection marker and a cloning site, but they carry one, or more recently two, cos sites derived from bacteriophage lambda. Depending on the particular aim of the experiment, broad host range cosmids, shuttle cosmids or 'mammalian' cosmids (linked to SV40 oriV and mammalian selection markers) are available. The loading capacity of cosmids varies depending on the size of the vector itself but usually lies around 40–45 kb. The cloning procedure involves the generation of two vector arms which are then joined to the foreign DNA. Selection against wild type cosmid DNA is simply done via size exclusion. Cosmids, therefore, always form colonies and not plaques. Also the clone density is much lower with around 10<sup>5</sup> – 10<sup>6</sup> <a href="Colony-forming_unit" title="Colony-forming unit">CFU</a> per μg of ligated DNA.

</p><p>After the construction of recombinant lambda or cosmid libraries the total DNA is transferred into an appropriate <i>E. coli</i> host via a technique called in vitro packaging. The necessary packaging extracts are derived from <i>E. coli</i> cI857 lysogens (red- gam- Sam and Dam (head assembly) and Eam (tail assembly) respectively). These extracts will recognize and package the recombinant molecules <i>in vitro</i>, generating either mature phage particles (lambda-based vectors) or recombinant plasmids contained in phage shells (cosmids). These differences are reflected in the different infection frequencies seen in favour of lambda-replacement vectors. This compensates for their slightly lower loading capacity. Phage libraries are also stored and screened more easily than cosmid libraries.

</p><p>Target DNA: the genomic DNA to be cloned has to be cut into the appropriate size range of restriction fragments. This is usually done by partial restriction followed by either size fractionation or dephosphorylation (using calf-intestine phosphatase) to avoid chromosome scrambling, i.e. the ligation of physically unlinked fragments.

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<div class="mw-heading mw-heading2"><h2 id="Examples_of_cosmid_vectors">Examples of cosmid vectors</h2></div>

<p>Commonly used vectors include "SuperCos 1"

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<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>

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</style><cite id="CITEREFHohnMurray1977" class="citation journal cs1">Hohn B, Murray K (August 1977). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC431522">"Packaging recombinant DNA molecules into bacteriophage particles in vitro"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>74</b> (8): <span class="nowrap">3259–</span>63. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1977PNAS...74.3259H">1977PNAS...74.3259H</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.74.8.3259">10.1073/pnas.74.8.3259</a></span>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC431522">431522</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/333431">333431</a>.</cite></span>

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<li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><cite id="CITEREFCollinsHohn1978" class="citation journal cs1">Collins J, Hohn B (September 1978). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC336088">"Cosmids: a type of plasmid gene-cloning vector that is packageable in vitro in bacteriophage lambda heads"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>75</b> (9): <span class="nowrap">4242–</span>6. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1978PNAS...75.4242C">1978PNAS...75.4242C</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.75.9.4242">10.1073/pnas.75.9.4242</a></span>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC336088">336088</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/360212">360212</a>.</cite></span>

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<li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.ndsu.edu/pubweb/~mcclean/plsc731/cloning/cloning4.htm">"Cosmids"</a>. Phillip McClean<span class="reference-accessdate">. Retrieved <span class="nowrap">April 4,</span> 2014</span>.</cite></span>

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<li id="cite_note-pmid6271486-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid6271486_4-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFCollins1981" class="citation journal cs1 cs1-prop-long-vol">Collins J (1981). "Instability of palindromic DNA in Escherichia coli". <i>Cold Spring Harbor Symposia on Quantitative Biology</i>. 45 Pt 1: <span class="nowrap">409–</span>16. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1101%2FSQB.1981.045.01.055">10.1101/SQB.1981.045.01.055</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/6271486">6271486</a>.</cite></span>

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<div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2></div>

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<ul><li>Stryer, Lubert (1995) <i>Biochemistry</i> 4th ed. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>0-7167-2009-4</bdi></li>

<li><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK6485/">Eurekah Biosciences Collection: Viruses, @NCBI</a></li></ul>

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</style><div id="Self-replicating_organic_structures216" style="font-size:114%;margin:0 4em"><a href="Self-replication" title="Self-replication">Self-replicating</a> organic structures</div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Life" title="Life">Cellular life</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Bacteria" title="Bacteria">Bacteria</a></li>

<li><a href="Archaea" title="Archaea">Archaea</a></li>

<li><a href="Eukaryote" title="Eukaryote">Eukaryota</a>

<ul><li><a href="Animal" title="Animal">Animalia</a></li>

<li><a href="Fungus" title="Fungus">Fungi</a></li>

<li><a href="Plant" title="Plant">Plantae</a></li>

<li><a href="Protist" title="Protist">Protista</a></li></ul></li>

<li><i><a href="Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i>

<ul><li><i><a href="Parakaryon" title="Parakaryon">Parakaryon</a></i></li>

<li><a href="Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="Virus" title="Virus">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a>

<ul><li><a href="Giant_virus" title="Giant virus">Giant virus</a></li></ul></li>

<li><a href="DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li>

<li><a href="Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li>

<li><a href="RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li>

<li><a href="RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li>

<li><a href="Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li>

<li><a href="DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral<br>agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li>

<li><i><a href="Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="Helper_virus" title="Helper virus">Helper-virus<br>dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;"><a href="Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li>ssRNA satellite virus</li>

<li>dsDNA satellite virus (<a href="Virophage" title="Virophage">Virophage</a>)</li>

<li>ssDNA satellite virus</li>

<li>ssDNA satellite</li>

<li>dsRNA satellite</li>

<li>ssRNA satellite (<a href="Virusoid" title="Virusoid">Virusoid</a>)</li>

<li>Satellite-like nucleic acids

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Prion" title="Prion">Mammalian prion</a></li>

<li><a href="Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Nucleic_acid" title="Nucleic acid">Nucleic acid</a><br>self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic<br>elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Mobilome" title="Mobilome">Mobilome</a>

<ul><li><a href="Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li>

<li><a href="Genomic_island" title="Genomic island">Genomic island</a></li></ul></li>

<li><a href="Transposable_element" title="Transposable element">Transposable element</a>

<ul><li><a href="Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li>

<li><a href="DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li>

<li><a href="Plasmid" title="Plasmid">Plasmid</a>

<ul><li><a href="Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li>

<li><a href="R-factor" class="mw-redirect" title="R-factor">Resistance</a></li>

<li>Degradative</li>

<li><a href="Virulence_factor" title="Virulence factor">Virulence</a>/<a href="Ti_plasmid" title="Ti plasmid">Ti</a></li>

<li>Cryptic</li></ul></li>

<li>

<ul><li><a href="Fosmid" title="Fosmid">Fosmid</a></li></ul></li>

<li><a href="Phagemid" title="Phagemid">Phagemid</a></li>

<li><a href="Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li>

<li><a href="Group_II_intron" title="Group II intron">Group II intron</a></li>

<li><a href="Retrozyme" title="Retrozyme">Retrozyme</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em">

<ul><li><a href="DNA_replication" title="DNA replication">DNA replication</a>

<ul><li><a href="RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li>

<li><a href="Chromosome" title="Chromosome">Chromosome</a>

<ul><li><a href="Linear_chromosome" title="Linear chromosome">Linear</a></li>

<li><a href="Circular_chromosome" title="Circular chromosome">Circular</a></li>

<li><a href="Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li>

<li><a href="Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li>

<li><a href="Genome" title="Genome">Genome</a>

<li><a href="Gene_duplication" title="Gene duplication">Gene duplication</a></li>

<li><a href="Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li>

<li><a href="Origin_of_replication" title="Origin of replication">Origin of replication</a>

<ul><li><a href="Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li>

<li><a href="Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a>

<ul><li><a href="Provirus" title="Provirus">Provirus</a></li>

<li><a href="Prophage" title="Prophage">Prophage</a></li>

<li><a href="Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li>

<li><a href="Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li>

<li><a href="Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a>

<ul><li><a href="Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li>

<li><a href="Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Mitochondrion" title="Mitochondrion">Mitochondrion</a>

<ul><li><a href="Mitosome" title="Mitosome">Mitosome</a></li>

<li><a href="Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li>

<li><a href="Plastid" title="Plastid">Plastid</a>

<ul><li><a href="Chloroplast" title="Chloroplast">Chloroplast</a></li>

<li><a href="Chromoplast" title="Chromoplast">Chromoplast</a></li>

<li><a href="Gerontoplast" title="Gerontoplast">Gerontoplast</a></li>

<li><a href="Leucoplast" title="Leucoplast">Leucoplast</a></li>

<li><a href="Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li>

<li><a href="Kappa_organism" title="Kappa organism">Kappa organism</a></li>

<li>Organs

<ul><li><a href="Bacteriome" title="Bacteriome">Bacteriome</a></li>

<li><a href="Trophosome" title="Trophosome">Trophosome</a></li></ul></li>

<li><a href="Nitroplast" title="Nitroplast">Nitroplast</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li>

<li><a href="Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li>

<ul><li><a href="Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li>

<li>†<a href="Protocell" title="Protocell">Protocell</a></li>

<li><a href="Coacervate" title="Coacervate">Coacervate</a></li>

<li><a href="Proteinoid" title="Proteinoid">Proteinoid</a></li>

<li><a href="Sulphobes" title="Sulphobes">Sulphobe</a></li>

<li>Research

<ul><li><a href="Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li>

<li><a href="Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Organism" title="Organism">Organism</a></li>

<ul><li><a href="Cell_division" title="Cell division">Cell division</a></li>

<li><a href="Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li>

<li><a href="Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li>

<li><a href="Synthetic_virology" title="Synthetic virology">Synthetic virus</a>

<ul><li><a href="Viral_vector" title="Viral vector">Viral vector</a></li>

<li><a href="Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li>

<li>?<a href="Nanobacterium" title="Nanobacterium">Nanobacterium</a></li>

<li>?<a href="Nanobe" title="Nanobe">Nanobe</a></li>

<li><a href="Cancer_cell" title="Cancer cell">Cancer cell</a>

<li><a href="Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li>

<li><a href="Virome" title="Virome">Virome</a></li></ul>

</div></td></tr></tbody></table></div>

<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Life,_non-cellular_life,_and_comparable_structures112" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Life,_non-cellular_life,_and_comparable_structures112" style="font-size:114%;margin:0 4em"><a href="Life" title="Life">Life</a>, <a href="Non-cellular_life" title="Non-cellular life">non-cellular life</a>, and comparable structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">"<a href="Prokaryote" title="Prokaryote">Prokaryota</a>"</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Archaea" title="Archaea">Archaea</a></li>

<li><a href="Bacteria" title="Bacteria">Bacteria</a>

<ul><li><a href="Mitochondrion" title="Mitochondrion">Mitochondrion</a></li>

<li><a href="Plastid" title="Plastid">Plastid</a></li></ul></li>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Eukaryote" title="Eukaryote">Eukaryota</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Animal" title="Animal">Animalia</a></li>

<li><a href="Fungus" title="Fungus">Fungi</a></li>

<li><a href="Plant" title="Plant">Plantae</a></li>

<li>"<a href="Protist" title="Protist">Protista</a>"</li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Parakaryon" title="Parakaryon">Parakaryon</a></i></li>

<li><a href="Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Virus" title="Virus">Virus</a>

<hr><a href="Viroid" title="Viroid">Viroid</a>

<hr><a href="Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Realms</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Adnaviria" title="Adnaviria">Adnaviria</a></i></li>

<li><i><a href="Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></i></li>

<li><i><a href="Monodnaviria" title="Monodnaviria">Monodnaviria</a></i></li>

<li><i><a href="Riboviria" title="Riboviria">Riboviria</a></i></li>

<li><i><a href="Ribozyviria" title="Ribozyviria">Ribozyviria</a></i></li>

<li><i><a href="Varidnaviria" title="Varidnaviria">Varidnaviria</a></i></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Unassigned</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Classes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><i><a href="Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></i></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Families</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></i></li>

<li><i><a href="Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></i></li>

<li><i><a href="Anelloviridae" title="Anelloviridae">Anelloviridae</a></i></li>

<li><i><a href="Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li>

<li><i><a href="Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></i></li>

<li><i><a href="Clavaviridae" title="Clavaviridae">Clavaviridae</a></i></li>

<li><i><a href="Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></i></li>

<li><i><a href="Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></i></li>

<li><i><a href="Globuloviridae" title="Globuloviridae">Globuloviridae</a></i></li>

<li><i><a href="Guttaviridae" title="Guttaviridae">Guttaviridae</a></i></li>

<li><i><a href="Halspiviridae" title="Halspiviridae">Halspiviridae</a></i></li>

<li><i><a href="Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></i></li>

<li><i><a href="Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></i></li>

<li><i><a href="Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></i></li>

<li><i><a href="Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></i></li>

<li><i><a href="Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li>

<li><i><a href="Spiraviridae" title="Spiraviridae">Spiraviridae</a></i></li>

<li><i><a href="Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></i></li>

<li><i><a href="Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></i></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genera</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></i></li>

<li><i><a href="Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></i></li></ul>

</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Nanobe" title="Nanobe">Nanobe</a> (?)</li>

<li><a href="Obelisk_(biology)" title="Obelisk (biology)">Obelisk</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Comparable<br>structures</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Bio-like_structure" title="Bio-like structure">Bio-like structure</a></li>

<li><a href="Cancer_cell" title="Cancer cell">Cancer cell</a>

<li><a href="Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li>

<li><a href="Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a></li>

<li><a href="Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li>

<li><a href="Fosmid" title="Fosmid">Fosmid</a></li>

<li><a href="Integrative_and_conjugative_element" title="Integrative and conjugative element">Integrative and conjugative element</a></li>

<li><a href="Jeewanu" title="Jeewanu">Jeewanu</a></li>

<li>"<i><a href="Nanobacterium" title="Nanobacterium">Nanobacterium</a></i>"</li>

<li><a href="Phagemid" title="Phagemid">Phagemid</a></li>

<li><a href="Plasmid" title="Plasmid">Plasmid</a></li>

<li><a href="Prion" title="Prion">Prion</a>

<ul><li><a href="Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul></li>

<li><a href="Proteinoid" title="Proteinoid">Proteinoid</a> <a href="Microparticle#Biological_protocells" title="Microparticle">microsphere</a></li>

<li>Retroelements not elsewhere classified

<ul><li><a href="Retron" title="Retron">Retron</a></li>

<li><a href="Diversity-generating_retroelement" title="Diversity-generating retroelement">Diversity-generating retroelement</a></li>

<li><a href="Telomerase_reverse_transcriptase" title="Telomerase reverse transcriptase">Telomerase reverse transcriptase</a></li>

<li><a href="Reverse_transcriptase-related_cellular_gene" class="mw-redirect" title="Reverse transcriptase-related cellular gene">Reverse transcriptase-related cellular gene</a></li></ul></li>

<li><a href="Ribozyme" title="Ribozyme">Ribozyme</a></li>

<li><a href="Spiegelman's_Monster" title="Spiegelman's Monster">Spiegelman's Monster</a></li>

<li><a href="Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li>

<li><a href="Transposable_element" title="Transposable element">Transposable element</a>

<ul><li><a href="Retroposon" title="Retroposon">Retroposon</a></li></ul></li>

<li><a href="Transpoviron" title="Transpoviron">Transpoviron</a></li>

<li><a href="Xenobot" title="Xenobot">Xenobot</a></li></ul>

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