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Crystallin - Wikipedia

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</div> </div> </div> </div> </nav> </div> </div> </div> <div class="vector-column-end"> <div class="vector-sticky-pinned-container"> <nav class="vector-page-tools-landmark" aria-label="Page tools"> <div id="vector-page-tools-pinned-container" class="vector-pinned-container"> </div> </nav> <nav class="vector-appearance-landmark" aria-label="Appearance"> <div id="vector-appearance-pinned-container" class="vector-pinned-container"> <div id="vector-appearance" class="vector-appearance vector-pinnable-element"> <div class="vector-pinnable-header vector-appearance-pinnable-header vector-pinnable-header-pinned" data-feature-name="appearance-pinned" data-pinnable-element-id="vector-appearance" data-pinned-container-id="vector-appearance-pinned-container" data-unpinned-container-id="vector-appearance-unpinned-container" > <div class="vector-pinnable-header-label">Appearance</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" 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div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Crystalline" class="mw-redirect" title="Crystalline">Crystalline</a>.</div> <p>In <a href="/wiki/Anatomy" title="Anatomy">anatomy</a>, a <b>crystallin</b> is a water-soluble structural <a href="/wiki/Protein" title="Protein">protein</a> found in the <a href="/wiki/Lens_(anatomy)" class="mw-redirect" title="Lens (anatomy)">lens</a> and the <a href="/wiki/Cornea" title="Cornea">cornea</a> of the <a href="/wiki/Eye" title="Eye">eye</a> accounting for the transparency of the structure.<sup id="cite_ref-pmid17997336_1-0" class="reference"><a href="#cite_note-pmid17997336-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> It has also been identified in other places such as the heart, and in aggressive breast cancer tumors.<sup id="cite_ref-heart-presence-1_2-0" class="reference"><a href="#cite_note-heart-presence-1-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-tumor-presence-1_3-0" class="reference"><a href="#cite_note-tumor-presence-1-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup> The physical origins of eye lens transparency and its relationship to <a href="/wiki/Cataract" title="Cataract">cataract</a> are an active area of research. <sup id="cite_ref-Crystallin_review_4-0" class="reference"><a href="#cite_note-Crystallin_review-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> Since it has been shown that lens injury may promote nerve regeneration,<sup id="cite_ref-nerve-regeneration-1_5-0" class="reference"><a href="#cite_note-nerve-regeneration-1-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> crystallin has been an area of neural research. So far, it has been demonstrated that crystallin β b2 (crybb2) may be a <a href="/wiki/Neurite" title="Neurite">neurite</a>-promoting factor.<sup id="cite_ref-neurite-promoting-1_6-0" class="reference"><a href="#cite_note-neurite-promoting-1-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=1" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The main function of crystallins at least in the <a href="/wiki/Lens_(anatomy)" class="mw-redirect" title="Lens (anatomy)">lens</a> of the eye is probably to increase the <a href="/wiki/Refractive_index" title="Refractive index">refractive index</a> while not obstructing light. However, this is not their only function. It has become clear that crystallins may have several <a href="/wiki/Metabolic" class="mw-redirect" title="Metabolic">metabolic</a> and regulatory functions, both within the lens and in other parts of the body.<sup id="cite_ref-pmid12790344_7-0" class="reference"><a href="#cite_note-pmid12790344-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> More proteins containing βγ-crystallin domains have now been characterized as calcium binding proteins with <a href="/wiki/Greek_key_(protein_structure)" class="mw-redirect" title="Greek key (protein structure)">Greek key</a> motif as a novel calcium-binding motif.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Enzyme_activity">Enzyme activity</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=2" title="Edit section: Enzyme activity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some crystallins are active <a href="/wiki/Enzymes" class="mw-redirect" title="Enzymes">enzymes</a>, while others lack activity but show <a href="/wiki/Sequence_homology" title="Sequence homology">homology</a> to other enzymes.<sup id="cite_ref-Enzymatic-activity-1_9-0" class="reference"><a href="#cite_note-Enzymatic-activity-1-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Enzymatic-activity-2_10-0" class="reference"><a href="#cite_note-Enzymatic-activity-2-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> The crystallins of different groups of organisms are related to a large number of different proteins, with those from birds and reptiles related to <a href="/wiki/Lactate_dehydrogenase" title="Lactate dehydrogenase">lactate dehydrogenase</a> and <a href="/wiki/Argininosuccinate_lyase" title="Argininosuccinate lyase">argininosuccinate lyase</a>, those of mammals to <a href="/wiki/Alcohol_dehydrogenase" title="Alcohol dehydrogenase">alcohol dehydrogenase</a> and <a href="/wiki/NADPH:quinone_reductase" title="NADPH:quinone reductase">quinone reductase</a>, and those of <a href="/wiki/Cephalopod" title="Cephalopod">cephalopods</a> to <a href="/wiki/Glutathione_S-transferase" title="Glutathione S-transferase">glutathione S-transferase</a> and <a href="/wiki/Aldehyde_dehydrogenase" title="Aldehyde dehydrogenase">aldehyde dehydrogenase</a>. Whether these crystallins are products of a fortuitous accident of evolution, in that these particular enzymes happened to be transparent and highly soluble, or whether these diverse enzymatic activities are part of the protective machinery of the lens, is an active research topic.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> The recruitment of protein that originally evolved with one function to serve a second, unrelated function is an example of an <a href="/wiki/Exaptation" title="Exaptation">exaptation</a>.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> </p> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:AlignmentHumanCrystallinABG.PNG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/AlignmentHumanCrystallinABG.PNG/200px-AlignmentHumanCrystallinABG.PNG" decoding="async" width="200" height="93" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/AlignmentHumanCrystallinABG.PNG/300px-AlignmentHumanCrystallinABG.PNG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7a/AlignmentHumanCrystallinABG.PNG/400px-AlignmentHumanCrystallinABG.PNG 2x" data-file-width="657" data-file-height="307" /></a><figcaption>An alignment of the human crystallin proteins alpha, beta, and gamma from Uniprot.</figcaption></figure> <div class="mw-heading mw-heading2"><h2 id="Classification">Classification</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=3" title="Edit section: Classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Crystallins from a vertebrate eye lens are classified into three main types: alpha, beta and gamma crystallins. These distinctions are based on the order in which they <a href="/wiki/Elution" title="Elution">elute</a> from a <a href="/wiki/Size-exclusion_chromatography" title="Size-exclusion chromatography">gel filtration chromatography</a> column. These are also called ubiquitous crystallins. Beta- and gamma-crystallins (such as <a href="/wiki/CRYGC" title="CRYGC">CRYGC</a>) are similar in sequence, structure and domains topology, and thus have been grouped together as a <a href="/wiki/Protein_superfamily" title="Protein superfamily">protein superfamily</a> called βγ-Crystallins. The α-crystallin family and βγ-crystallins compose the major family of proteins present in the crystalline lens. They occur in all vertebrate classes (though gamma-crystallins are low or absent in avian lenses); and delta-crystallin is found exclusively in reptiles and birds.<sup id="cite_ref-PUB00005345_13-0" class="reference"><a href="#cite_note-PUB00005345-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-PUB00003917_14-0" class="reference"><a href="#cite_note-PUB00003917-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup> </p><p>In addition to these crystallins there are other <a href="/wiki/Taxon" title="Taxon">taxon</a>-specific crystallins which are only found in the lens of some organisms; these include delta, epsilon, tau, and iota-crystallins. For example, alpha, beta, and delta crystallins are found in avian and reptilian lenses, and the alpha, beta, and gamma families are found in the lenses of all other vertebrates. </p> <div class="mw-heading mw-heading3"><h3 id="Alpha-crystallin">Alpha-crystallin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=4" title="Edit section: Alpha-crystallin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Protein family</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">Alpha crystallin A chain, N terminal</th></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd; color:inherit;">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;">Symbol</th><td class="infobox-data" style="background-color: #eee; color:inherit;">Crystallin</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Pfam" title="Pfam">Pfam</a></th><td class="infobox-data pfam" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF00525">PF00525</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/InterPro" title="InterPro">InterPro</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/IPR003090">IPR003090</a></td></tr><tr><td colspan="2" class="infobox-full-data" style="background-color: #eee; color:inherit;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546" /><table class="infobox mw-collapsible mw-collapsed" style="float:none; clear:none; margin:0; border-width:0; border-collapse:collapse; text-align:left; width:100%"><tbody><tr><th colspan="2" class="infobox-header" style="background-color: #ddd; color:inherit;">Available protein structures:</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Pfam" title="Pfam">Pfam</a> &#160;</th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="http://pfam.xfam.org/family/PF00525?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF00525">ECOD</a> &#160;</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a></th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.rcsb.org/search?q=rcsb_polymer_entity_annotation.annotation_id:PF00525%20AND%20rcsb_polymer_entity_annotation.type:Pfam">RCSB PDB</a>; <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/entry/search/index?pfam_accession:PF00525">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&amp;other_db_field=PF00525">PDBj</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/PDBsum" title="PDBsum">PDBsum</a></th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/thornton-srv/databases/cgi-bin/pdbsum/GetPfamStr.pl?pfam_id=PF00525">structure summary</a></td></tr></tbody></table></td></tr></tbody></table> <p>Alpha-crystallin occurs as large aggregates, comprising two types of related subunits (A and B) that are highly similar to the small (15-30kDa) heat shock proteins (<a href="/wiki/SHsp" class="mw-redirect" title="SHsp">sHsps</a>), particularly in their C-terminal halves. The relationship between these families is one of classic gene duplication and divergence, from the small HSP family, allowing adaptation to novel functions. Divergence probably occurred prior to evolution of the eye lens, alpha-crystallin being found in small amounts in tissues outside the lens.<sup id="cite_ref-PUB00005345_13-1" class="reference"><a href="#cite_note-PUB00005345-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>Alpha-crystallin has <a href="/wiki/Chaperone_(protein)" title="Chaperone (protein)">chaperone</a>-like properties including the ability to prevent the precipitation of denatured proteins and to increase cellular tolerance to stress.<sup id="cite_ref-PUB00034659_15-0" class="reference"><a href="#cite_note-PUB00034659-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> It has been suggested that these functions are important for the maintenance of lens transparency and the prevention of <a href="/wiki/Cataract" title="Cataract">cataracts</a>.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup> This is supported by the observation that alpha-crystallin mutations show an association with cataract formation. </p><p>The N-terminal domain of alpha-crystallin is not necessary for dimerisation or chaperone activity, but appears to be required for the formation of higher order aggregates.<sup id="cite_ref-PUB00034662_17-0" class="reference"><a href="#cite_note-PUB00034662-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-PUB00034663_18-0" class="reference"><a href="#cite_note-PUB00034663-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Beta_and_gamma_crystallin">Beta and gamma crystallin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=5" title="Edit section: Beta and gamma crystallin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Protein family</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546" /><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">Beta/Gamma crystallin</th></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd; color:inherit;">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;">Symbol</th><td class="infobox-data" style="background-color: #eee; color:inherit;">Crystall</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Pfam" title="Pfam">Pfam</a></th><td class="infobox-data pfam" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF00030">PF00030</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/InterPro" title="InterPro">InterPro</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/IPR001064">IPR001064</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/PROSITE" title="PROSITE">PROSITE</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://prosite.expasy.org/PDOC00197">PDOC00197</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Structural_Classification_of_Proteins" class="mw-redirect" title="Structural Classification of Proteins">SCOP2</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="http://scop2.mrc-lmb.cam.ac.uk/search?t=txt;q=4gcr">4gcr</a> / <a rel="nofollow" class="external text" href="https://scop.berkeley.edu/pdb/code=4gcr">SCOPe</a> / <a rel="nofollow" class="external text" href="http://supfam.org/SUPERFAMILY/cgi-bin/search.cgi?search_field=4gcr">SUPFAM</a></td></tr><tr><td colspan="2" class="infobox-full-data" style="background-color: #eee; color:inherit;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546" /><table class="infobox mw-collapsible mw-collapsed" style="float:none; clear:none; margin:0; border-width:0; border-collapse:collapse; text-align:left; width:100%"><tbody><tr><th colspan="2" class="infobox-header" style="background-color: #ddd; color:inherit;">Available protein structures:</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Pfam" title="Pfam">Pfam</a> &#160;</th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="http://pfam.xfam.org/family/PF00030?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF00030">ECOD</a> &#160;</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a></th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.rcsb.org/search?q=rcsb_polymer_entity_annotation.annotation_id:PF00030%20AND%20rcsb_polymer_entity_annotation.type:Pfam">RCSB PDB</a>; <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/entry/search/index?pfam_accession:PF00030">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&amp;other_db_field=PF00030">PDBj</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/PDBsum" title="PDBsum">PDBsum</a></th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/thornton-srv/databases/cgi-bin/pdbsum/GetPfamStr.pl?pfam_id=PF00030">structure summary</a></td></tr></tbody></table></td></tr></tbody></table> <p>Beta- and gamma- crystallin form a separate family.<sup id="cite_ref-PUB00003409_19-0" class="reference"><a href="#cite_note-PUB00003409-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-PUB00004933_20-0" class="reference"><a href="#cite_note-PUB00004933-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> Structurally, beta and gamma crystallins are composed of two similar domains which, in turn, are each composed of two similar <a href="/wiki/Structural_motif" title="Structural motif">motifs</a> with the two domains connected by a short connecting <a href="/wiki/Peptide" title="Peptide">peptide</a>. Each motif, which is about forty amino acid residues long, is folded in a distinctive <a href="/wiki/Beta_sheet#Greek_key_motif" title="Beta sheet">Greek key pattern</a>. However, beta crystallin is an <a href="/wiki/Oligomer" title="Oligomer">oligomer</a>, composed of a complex group of molecules, whereas gamma crystallin is a simpler <a href="/wiki/Monomer" title="Monomer">monomer</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> <sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Cleanup plainlinks metadata ambox ambox-style ambox-Cleanup" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section may <b>require <a href="/wiki/Wikipedia:Cleanup" title="Wikipedia:Cleanup">cleanup</a></b> to meet Wikipedia's <a href="/wiki/Wikipedia:Manual_of_Style" title="Wikipedia:Manual of Style">quality standards</a>. The specific problem is: <b>Lots of duplicate entries in the table. Please regenerate the table using only the "reviewed" entries.</b><span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Crystallin" title="Special:EditPage/Crystallin">improve this section</a> if you can.</span> <span class="date-container"><i>(<span class="date">March 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <table class="wikitable collapsible" width="250px"> <caption>List of Human Crystallins <sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> </caption> <tbody><tr> <th>UniProt Entry name</th> <th>Alternate Gene names</th> <th>Length </th></tr> <tr> <td>AIM1L_HUMAN</td> <td>AIM1L CRYBG2</td> <td>616 </td></tr> <tr> <td>AIM1_HUMAN</td> <td>AIM1 CRYBG1</td> <td>1723 </td></tr> <tr> <td>ARLY_HUMAN</td> <td>ASL</td> <td>464 </td></tr> <tr> <td>CRBA1_HUMAN</td> <td>CRYBA1 CRYB1</td> <td>215 </td></tr> <tr> <td>CRBA2_HUMAN</td> <td>CRYBA2</td> <td>197 </td></tr> <tr> <td>CRBB1_HUMAN</td> <td>CRYBB1</td> <td>252 </td></tr> <tr> <td>CRBA4_HUMAN</td> <td>CRYBA4</td> <td>196 </td></tr> <tr> <td>CRBB2_HUMAN</td> <td>CRYBB2 CRYB2 CRYB2A</td> <td>205 </td></tr> <tr> <td>CRBB3_HUMAN</td> <td>CRYBB3 CRYB3</td> <td>211 </td></tr> <tr> <td>CRBG3_HUMAN</td> <td>CRYBG3</td> <td>1022 </td></tr> <tr> <td>CRBS_HUMAN</td> <td>CRYGS CRYG8</td> <td>178 </td></tr> <tr> <td>CRGA_HUMAN</td> <td>CRYGA CRYG1</td> <td>174 </td></tr> <tr> <td>CRGC_HUMAN</td> <td>CRYGC CRYG3</td> <td>174 </td></tr> <tr> <td>CRGB_HUMAN</td> <td>CRYGB CRYG2</td> <td>175 </td></tr> <tr> <td>CRGN_HUMAN</td> <td>CRYGN</td> <td>182 </td></tr> <tr> <td>CRGD_HUMAN</td> <td>CRYGD CRYG4</td> <td>174 </td></tr> <tr> <td>CRYAA_HUMAN</td> <td>CRYAA CRYA1 HSPB4</td> <td>173 </td></tr> <tr> <td>CRYAB_HUMAN</td> <td>CRYAB CRYA2</td> <td>175 </td></tr> <tr> <td>CRYL1_HUMAN</td> <td>CRYL1 CRY</td> <td>319 </td></tr> <tr> <td>CRYM_HUMAN</td> <td>CRYM THBP</td> <td>314 </td></tr> <tr> <td>HSPB2_HUMAN</td> <td>HSPB2</td> <td>182 </td></tr> <tr> <td>HSPB3_HUMAN</td> <td>HSPB3 HSP27 HSPL27</td> <td>150 </td></tr> <tr> <td>HSPB8_HUMAN</td> <td>HSPB8 CRYAC E2IG1 HSP22 PP1629</td> <td>196 </td></tr> <tr> <td>HSPB7_HUMAN</td> <td>HSPB7 CVHSP</td> <td>170 </td></tr> <tr> <td>HSPB9_HUMAN</td> <td>HSPB9</td> <td>159 </td></tr> <tr> <td>HSPB1_HUMAN</td> <td>HSPB1 HSP27 HSP28</td> <td>205 </td></tr> <tr> <td>HSPB6_HUMAN</td> <td>HSPB6</td> <td>160 </td></tr> <tr> <td>IFT25_HUMAN</td> <td>HSPB11 C1orf41 IFT25 HSPC034</td> <td>144 </td></tr> <tr> <td>MAF_HUMAN</td> <td>MAF</td> <td>373 </td></tr> <tr> <td>ODFP1_HUMAN</td> <td>ODF1 ODFP</td> <td>250 </td></tr> <tr> <td>QORL1_HUMAN</td> <td>CRYZL1 4P11</td> <td>349 </td></tr> <tr> <td>QOR_HUMAN</td> <td>CRYZ</td> <td>329 </td></tr> <tr> <td>TITIN_HUMAN</td> <td>TTN</td> <td>34350 </td></tr> <tr> <td>ZEB1_HUMAN</td> <td>ZEB1 AREB6 TCF8</td> <td>1124 </td></tr> <tr> <td>Q9UFA7_HUMAN</td> <td>DKFZp434A0627 CRYGS hCG_16149</td> <td>120 </td></tr> <tr> <td>B4DU04_HUMAN</td> <td>AIM1 hCG_33516</td> <td>542 </td></tr> <tr> <td>A8KAH6_HUMAN</td> <td>HSPB2 hCG_39461</td> <td>182 </td></tr> <tr> <td>Q6ICS9_HUMAN</td> <td>HSPB3 hCG_1736006</td> <td>150 </td></tr> <tr> <td>Q68DG0_HUMAN</td> <td>DKFZp779D0968 HSPB7</td> <td>174 </td></tr> <tr> <td>Q8N241_HUMAN</td> <td>HSPB7 hCG_23506</td> <td>245 </td></tr> <tr> <td>B4DLE8_HUMAN</td> <td>CRYBG3</td> <td>1365 </td></tr> <tr> <td>C3VMY8_HUMAN</td> <td>CRYAB</td> <td>175 </td></tr> <tr> <td>R4UMM2_HUMAN</td> <td>CRYBB2</td> <td>205 </td></tr> <tr> <td>B3KQL3_HUMAN</td> <td></td> <td>119 </td></tr> <tr> <td>Q24JT5_HUMAN</td> <td>CRYGA</td> <td>105 </td></tr> <tr> <td>V9HWB6_HUMAN</td> <td>HEL55</td> <td>160 </td></tr> <tr> <td>B4DNC2_HUMAN</td> <td></td> <td>196 </td></tr> <tr> <td>V9HW27_HUMAN</td> <td>HEL-S-101</td> <td>175 </td></tr> <tr> <td>H0YCW8_HUMAN</td> <td>CRYAB</td> <td>106 </td></tr> <tr> <td>E9PHE4_HUMAN</td> <td>CRYAA</td> <td>136 </td></tr> <tr> <td>E9PNH7_HUMAN</td> <td>CRYAB</td> <td>106 </td></tr> <tr> <td>E7EWH7_HUMAN</td> <td>CRYAA</td> <td>153 </td></tr> <tr> <td>B4DL87_HUMAN</td> <td></td> <td>170 </td></tr> <tr> <td>V9HW43_HUMAN</td> <td>HEL-S-102</td> <td>205 </td></tr> <tr> <td>E9PR44_HUMAN</td> <td>CRYAB</td> <td>174 </td></tr> <tr> <td>Q8IVN0_HUMAN</td> <td></td> <td>86 </td></tr> <tr> <td>B7ZAH2_HUMAN</td> <td></td> <td>542 </td></tr> <tr> <td>C9J5A3_HUMAN</td> <td>HSPB7</td> <td>124 </td></tr> <tr> <td>E9PRS4_HUMAN</td> <td>CRYAB</td> <td>69 </td></tr> <tr> <td>K7EP04_HUMAN</td> <td>HSPB6</td> <td>137 </td></tr> <tr> <td>I3L3Y1_HUMAN</td> <td>CRYM</td> <td>97 </td></tr> <tr> <td>H0YG30_HUMAN</td> <td>HSPB8</td> <td>152 </td></tr> <tr> <td>H9KVC2_HUMAN</td> <td>CRYM</td> <td>272 </td></tr> <tr> <td>E9PS12_HUMAN</td> <td>CRYAB</td> <td>77 </td></tr> <tr> <td>E9PIR9_HUMAN</td> <td>AIM1L</td> <td>787 </td></tr> <tr> <td>B4DUL6_HUMAN</td> <td></td> <td>80 </td></tr> <tr> <td>I3NI53_HUMAN</td> <td>CRYM</td> <td>140 </td></tr> <tr> <td>Q9NTH7_HUMAN</td> <td>DKFZp434L1713</td> <td>264 </td></tr> <tr> <td>J3KQW1_HUMAN</td> <td>AIM1L</td> <td>296 </td></tr> <tr> <td>Q96QW7_HUMAN</td> <td>AIM1</td> <td>316 </td></tr> <tr> <td>I3L2W5_HUMAN</td> <td>CRYM</td> <td>165 </td></tr> <tr> <td>B1AHR5_HUMAN</td> <td>CRYBB3</td> <td>113 </td></tr> <tr> <td>B4DLI1_HUMAN</td> <td></td> <td>403 </td></tr> <tr> <td>I3L325_HUMAN</td> <td>CRYM</td> <td>241 </td></tr> <tr> <td>Q7Z3C1_HUMAN</td> <td>DKFZp686A14192</td> <td>191 </td></tr> <tr> <td>B4DWM9_HUMAN</td> <td></td> <td>154 </td></tr> <tr> <td>Q71V83_HUMAN</td> <td>CRYAA</td> <td>69 </td></tr> <tr> <td>Q6P5P8_HUMAN</td> <td>AIM1</td> <td>326 </td></tr> <tr> <td>C9JDH2_HUMAN</td> <td>CRYBA2</td> <td>129 </td></tr> <tr> <td>B4DIA6_HUMAN</td> <td></td> <td>155 </td></tr> <tr> <td>Q13684_HUMAN</td> <td></td> <td>56 </td></tr> <tr> <td>F8WE04_HUMAN</td> <td>HSPB1</td> <td>186 </td></tr> <tr> <td>J3QRT1_HUMAN</td> <td>CRYBA1</td> <td>75 </td></tr> <tr> <td>E9PRA8_HUMAN</td> <td>CRYAB</td> <td>155 </td></tr> <tr> <td>E9PJL7_HUMAN</td> <td>CRYAB</td> <td>130 </td></tr> <tr> <td>C9J5N2_HUMAN</td> <td>CRYBG3</td> <td>229 </td></tr> <tr> <td>I3L3J9_HUMAN</td> <td>CRYM</td> <td>26 </td></tr> <tr> <td>C9J659_HUMAN</td> <td>CRYBG3</td> <td>131 </td></tr> <tr> <td>D3YTC6_HUMAN</td> <td>HSPB7</td> <td>165 </td></tr> </tbody></table> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=6" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-pmid17997336-1"><span 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Biol</i>. <b>51</b> (1): <span class="nowrap">47–</span>76. <a href="/wiki/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/10.1016%2F0079-6107%2888%2990010-7">10.1016/0079-6107(88)90010-7</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/3064189">3064189</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Prog.+Biophys.+Mol.+Biol.&amp;rft.atitle=The+evolution+of+lenticular+proteins%3A+the+beta-+and+gamma-crystallin+super+gene+family&amp;rft.volume=51&amp;rft.issue=1&amp;rft.pages=%3Cspan+class%3D%22nowrap%22%3E47-%3C%2Fspan%3E76&amp;rft.date=1988&amp;rft_id=info%3Adoi%2F10.1016%2F0079-6107%2888%2990010-7&amp;rft_id=info%3Apmid%2F3064189&amp;rft.aulast=Schoenmakers&amp;rft.aufirst=JG&amp;rft.au=Lubsen%2C+NH&amp;rft.au=Aarts%2C+HJ&amp;rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252F0079-6107%252888%252990010-7&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ACrystallin" class="Z3988"></span></span> </li> <li id="cite_note-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-21">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFNathaniel_Knox_CartwrightPetros_Carvounis2005" class="citation book cs1">Nathaniel Knox Cartwright; Petros Carvounis (2005). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=5KslZLOs58gC&amp;pg=PA80"><i>Short answer questions for the MRCOphth, Part 1</i></a>. Radcliffe Publishing. p.&#160;80. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/wiki/Special:BookSources/9781857758849" title="Special:BookSources/9781857758849"><bdi>9781857758849</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Short+answer+questions+for+the+MRCOphth%2C+Part+1&amp;rft.pages=80&amp;rft.pub=Radcliffe+Publishing&amp;rft.date=2005&amp;rft.isbn=9781857758849&amp;rft.au=Nathaniel+Knox+Cartwright&amp;rft.au=Petros+Carvounis&amp;rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D5KslZLOs58gC%26pg%3DPA80&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ACrystallin" class="Z3988"></span></span> </li> <li id="cite_note-22"><span class="mw-cite-backlink"><b><a href="#cite_ref-22">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFGhoshChauhan2019" class="citation cs2">Ghosh, Kalyan Sundar; Chauhan, Priyanka (2019), "Crystallins and Their Complexes", <a rel="nofollow" class="external text" href="https://doi.org/10.1007/978-3-030-28151-9_14"><i>Macromolecular Protein Complexes II: Structure and Function</i></a>, Subcellular Biochemistry, vol.&#160;93, Springer International Publishing, pp.&#160;<span class="nowrap">439–</span>460, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-3-030-28151-9_14">10.1007/978-3-030-28151-9_14</a>, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/wiki/Special:BookSources/978-3-030-28151-9" title="Special:BookSources/978-3-030-28151-9"><bdi>978-3-030-28151-9</bdi></a>, <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/31939160">31939160</a></cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=bookitem&amp;rft.atitle=Crystallins+and+Their+Complexes&amp;rft.btitle=Macromolecular+Protein+Complexes+II%3A+Structure+and+Function&amp;rft.series=Subcellular+Biochemistry&amp;rft.pages=%3Cspan+class%3D%22nowrap%22%3E439-%3C%2Fspan%3E460&amp;rft.pub=Springer+International+Publishing&amp;rft.date=2019&amp;rft_id=info%3Apmid%2F31939160&amp;rft_id=info%3Adoi%2F10.1007%2F978-3-030-28151-9_14&amp;rft.isbn=978-3-030-28151-9&amp;rft.aulast=Ghosh&amp;rft.aufirst=Kalyan+Sundar&amp;rft.au=Chauhan%2C+Priyanka&amp;rft_id=https%3A%2F%2Fdoi.org%2F10.1007%2F978-3-030-28151-9_14&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ACrystallin" class="Z3988"></span></span> </li> <li id="cite_note-23"><span class="mw-cite-backlink"><b><a href="#cite_ref-23">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.uniprot.org/">"Uniprot"</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=unknown&amp;rft.btitle=Uniprot&amp;rft_id=https%3A%2F%2Fwww.uniprot.org%2F&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ACrystallin" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=7" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFGraw_J1997" class="citation journal cs1">Graw J (1997). "The crystallins: Genes, proteins and diseases". <i>Biological Chemistry</i>. <b>378</b> (11): <span class="nowrap">1331–</span>1348. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1515%2Fbchm.1997.378.11.1299">10.1515/bchm.1997.378.11.1299</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/9426193">9426193</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Biological+Chemistry&amp;rft.atitle=The+crystallins%3A+Genes%2C+proteins+and+diseases&amp;rft.volume=378&amp;rft.issue=11&amp;rft.pages=%3Cspan+class%3D%22nowrap%22%3E1331-%3C%2Fspan%3E1348&amp;rft.date=1997&amp;rft_id=info%3Adoi%2F10.1515%2Fbchm.1997.378.11.1299&amp;rft_id=info%3Apmid%2F9426193&amp;rft.au=Graw+J&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ACrystallin" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Crystallin&amp;action=edit&amp;section=8" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Crystallins">Crystallins</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li> <li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=alpha-Crystallins">alpha-Crystallins</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li> <li><a rel="nofollow" class="external text" href="http://sirtophamhatt.cryst.bbk.ac.uk/bca/CNews/1998/Dec98/eyeln.html">Lens Crystallin Crystal Structures</a><sup class="noprint Inline-Template"><span style="white-space: nowrap;">&#91;<i><a href="/wiki/Wikipedia:Link_rot" title="Wikipedia:Link rot"><span title="&#160;Dead link tagged August 2017">permanent dead link</span></a></i><span style="visibility:hidden; color:transparent; padding-left:2px">&#8205;</span>&#93;</span></sup> by Christine Slingsby, Birkbeck College</li> <li><a rel="nofollow" class="external text" href="http://www.rcsb.org/pdb/101/motm.do?momID=127">Crystallins: Molecule of the Month</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20151025082725/http://www.rcsb.org/pdb/101/motm.do?momID=127">Archived</a> 2015-10-25 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a>, by David Goodsell, RCSB Protein Data Bank</li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol 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navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Rhodopsin" title="Rhodopsin">Rhodopsin</a></li> <li><a href="/wiki/Cone_opsin" class="mw-redirect" title="Cone opsin">Cone opsins</a> <ul><li><a href="/wiki/OPN1LW" title="OPN1LW">OPN1LW</a></li> <li><a href="/wiki/OPN1MW" title="OPN1MW">OPN1MW</a>/<a href="/wiki/OPN1MW2" title="OPN1MW2">OPN1MW2</a></li> <li><a href="/wiki/OPN1SW" title="OPN1SW">OPN1SW</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Opsin#Novel_type_2_opsin_groups" title="Opsin">nonvisual</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="/wiki/Melanopsin" title="Melanopsin">Melanopsin</a></li> <li><a href="/wiki/OPN3" title="OPN3">OPN3</a></li> <li><a href="/wiki/OPN5" title="OPN5">OPN5</a></li> <li><a href="/wiki/RRH" title="RRH">RRH</a></li> <li><a href="/wiki/RGR_(gene)" class="mw-redirect" title="RGR (gene)">RGR</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a class="mw-selflink selflink">Crystallin</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="/wiki/Alpha_crystallin" class="mw-redirect" title="Alpha crystallin">Alpha</a> (<a href="/wiki/CRYAA" title="CRYAA">A</a></li> <li><a href="/wiki/CRYAB" title="CRYAB">B</a>)</li> <li><a href="/wiki/Beta_gamma_crystallin" class="mw-redirect" title="Beta gamma crystallin">Beta</a> (<a href="/wiki/Crystallin,_beta_A1" title="Crystallin, beta A1">A1</a></li> <li><a href="/w/index.php?title=CRYBA2&amp;action=edit&amp;redlink=1" class="new" title="CRYBA2 (page does not exist)">A2</a></li> <li><a href="/wiki/CRYBA4" title="CRYBA4">A4</a></li> <li><a href="/wiki/CRYBB1" title="CRYBB1">B1</a></li> <li><a href="/wiki/CRYBB2" title="CRYBB2">B2</a></li> <li><a href="/wiki/CRYBB3" title="CRYBB3">B3</a>)</li> <li><a href="/wiki/Beta_gamma_crystallin" class="mw-redirect" title="Beta gamma crystallin">Gamma</a> (<a href="/wiki/CRYGA" title="CRYGA">A</a></li> <li><a href="/wiki/CRYGB" title="CRYGB">B</a></li> <li><a href="/wiki/CRYGC" title="CRYGC">C</a></li> <li><a href="/wiki/Crystallin,_gamma_D" title="Crystallin, gamma D">D</a></li> <li><a href="/w/index.php?title=CRYGN&amp;action=edit&amp;redlink=1" class="new" title="CRYGN (page does not exist)">N</a></li> <li><a href="/wiki/CRYGS" title="CRYGS">S</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Other</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="/wiki/Arrestin" title="Arrestin">Arrestin</a></li> <li><a href="/wiki/Guanylate_cyclase_activator" title="Guanylate cyclase activator">Guanylate cyclase activator</a></li> <li><a href="/wiki/Recoverin" title="Recoverin">Recoverin</a></li> <li><a href="/wiki/Rhodopsin_kinase" title="Rhodopsin kinase">Rhodopsin kinase</a></li></ul> </div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.eqiad.main‐8669bc5c8‐4f76c Cached time: 20250318155414 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 0.468 seconds Real time usage: 0.636 seconds Preprocessor visited node count: 3033/1000000 Post‐expand include size: 120764/2097152 bytes Template argument size: 5249/2097152 bytes Highest expansion depth: 17/100 Expensive parser function count: 7/500 Unstrip recursion depth: 1/20 Unstrip post‐expand size: 108450/5000000 bytes Lua time usage: 0.287/10.000 seconds Lua memory usage: 6505043/52428800 bytes Number of Wikibase entities loaded: 0/400 --> <!-- Transclusion expansion time 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