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<div class="page-heading"> <h1 id="firstHeading" class="firstHeading mw-first-heading">Cellular differentiation</h1> <div class="tagline"></div> </div> <ul id="p-associated-pages" class="minerva__tab-container"> <li class="minerva__tab selected"> <a class="minerva__tab-text" href="/wiki/Cellular_differentiation" rel="" data-event-name="tabs.subject">Article</a> </li> <li class="minerva__tab "> <a class="minerva__tab-text" href="/wiki/Talk:Cellular_differentiation" rel="discussion" data-event-name="tabs.talk">Talk</a> </li> </ul> <nav class="page-actions-menu"> <ul id="p-views" class="page-actions-menu__list"> <li id="language-selector" class="page-actions-menu__list-item"> <a role="button" href="#p-lang" data-mw="interface" data-event-name="menu.languages" title="Language" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet language-selector"> Language </a> </li> <li id="page-actions-watch" class="page-actions-menu__list-item"> <a role="button" id="ca-watch" href="/w/index.php?title=Special:UserLogin&returnto=Cellular+differentiation" data-event-name="menu.watch" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet menu__item--page-actions-watch"> Watch </a> </li> <li id="page-actions-edit" class="page-actions-menu__list-item"> <a role="button" id="ca-edit" href="/w/index.php?title=Cellular_differentiation&action=edit" data-event-name="menu.edit" data-mw="interface" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet edit-page menu__item--page-actions-edit"> Edit </a> </li> </ul> </nav>  <div id="mw-content-subtitle">(Redirected from <a href="/w/index.php?title=Cell_differentiation&redirect=no" class="mw-redirect" title="Cell differentiation">Cell differentiation</a>)</div> </div> <div id="bodyContent" class="content"> <div id="mw-content-text" class="mw-body-content"><script>function mfTempOpenSection(id){var block=document.getElementById("mf-section-"+id);block.className+=" open-block";block.previousSibling.className+=" open-block";}</script><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><section class="mf-section-0" id="mf-section-0"> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output 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">"Cell differentiation" redirects here. For the journal, see <a href="/wiki/Cell_Differentiation_(journal)" class="mw-redirect" title="Cell Differentiation (journal)">Cell Differentiation (journal)</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Cell_division" title="Cell division">Cell division</a>.</div> Cellular differentiation is the process in which a <a href="/wiki/Stem_cell" title="Stem cell">stem cell</a> changes from one type to a differentiated one.<a href="#cite_note-2">[2]</a><a href="#cite_note-3">[3]</a> Usually, the cell changes to a more specialized type. Differentiation happens multiple times during the development of a <a href="/wiki/Multicellular_organism" title="Multicellular organism">multicellular organism</a> as it changes from a simple <a href="/wiki/Zygote" title="Zygote">zygote</a> to a complex system of <a href="/wiki/Tissue_(biology)" title="Tissue (biology)">tissues</a> and cell types. Differentiation continues in adulthood as <a href="/wiki/Adult_stem_cell" title="Adult stem cell">adult stem cells</a> divide and create fully differentiated <a href="/wiki/Cell_division" title="Cell division">daughter cells</a> during tissue repair and during normal cell turnover. Some differentiation occurs in response to <a href="/wiki/Antigen" title="Antigen">antigen</a> exposure. Differentiation dramatically changes a cell's size, shape, <a href="/wiki/Membrane_potential" title="Membrane potential">membrane potential</a>, <a href="/wiki/Metabolism" title="Metabolism">metabolic activity</a>, and responsiveness to signals. These changes are largely due to highly controlled modifications in <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> and are the study of <a href="/wiki/Epigenetics" title="Epigenetics">epigenetics</a>. With a few exceptions, cellular differentiation almost never involves a change in the <a href="/wiki/DNA" title="DNA">DNA</a> sequence itself. Metabolic composition, however, gets dramatically altered<a href="#cite_note-4">[4]</a> where stem cells are characterized by abundant metabolites with highly unsaturated structures whose levels decrease upon differentiation. Thus, different cells can have very different physical characteristics despite having the same <a href="/wiki/Genome" title="Genome">genome</a>. <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Final_stem_cell_differentiation_(1).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Final_stem_cell_differentiation_%281%29.svg/330px-Final_stem_cell_differentiation_%281%29.svg.png" decoding="async" width="330" height="233" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Final_stem_cell_differentiation_%281%29.svg/495px-Final_stem_cell_differentiation_%281%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Final_stem_cell_differentiation_%281%29.svg/660px-Final_stem_cell_differentiation_%281%29.svg.png 2x" data-file-width="512" data-file-height="362"></a><figcaption><a href="/wiki/Stem_cell" title="Stem cell">Stem cell</a> differentiation into various animal tissue types</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Stimuli.pdf" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9b/Stimuli.pdf/page1-220px-Stimuli.pdf.jpg" decoding="async" width="220" height="214" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9b/Stimuli.pdf/page1-330px-Stimuli.pdf.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9b/Stimuli.pdf/page1-440px-Stimuli.pdf.jpg 2x" data-file-width="675" data-file-height="658"></a><figcaption>Cell-count distribution featuring cellular differentiation for three types of cells (progenitor <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle z}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>z</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle z}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/bf368e72c009decd9b6686ee84a375632e11de98" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:1.088ex; height:1.676ex;" alt="{\displaystyle z}">, osteoblast <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle y}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>y</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle y}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/b8a6208ec717213d4317e666f1ae872e00620a0d" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:1.155ex; height:2.009ex;" alt="{\displaystyle y}">, and chondrocyte <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle x}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>x</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle x}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/87f9e315fd7e2ba406057a97300593c4802b53e4" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:1.33ex; height:1.676ex;" alt="{\displaystyle x}">) exposed to pro-osteoblast stimulus.<a href="#cite_note-CME-1">[1]</a></figcaption></figure> A specialized type of differentiation, known as <a href="/wiki/Terminal_differentiation" class="mw-redirect" title="Terminal differentiation">terminal differentiation</a>, is of importance in some tissues, including vertebrate <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a>, <a href="/wiki/Striated_muscle" class="mw-redirect" title="Striated muscle">striated muscle</a>, <a href="/wiki/Epidermis" title="Epidermis">epidermis</a> and gut. During terminal differentiation, a precursor cell formerly capable of cell division permanently leaves the cell cycle, dismantles the cell cycle machinery and often expresses a range of genes characteristic of the cell's final function (e.g. <a href="/wiki/Myosin" title="Myosin">myosin</a> and <a href="/wiki/Actin" title="Actin">actin</a> for a muscle cell). Differentiation may continue to occur after terminal differentiation if the capacity and functions of the cell undergo further changes. Among dividing cells, there are multiple levels of <a href="/wiki/Cell_potency" title="Cell potency">cell potency</a>, which is the cell's ability to differentiate into other cell types. A greater potency indicates a larger number of cell types that can be derived. A cell that can differentiate into all cell types, including the placental tissue, is known as <a href="/wiki/Totipotent" class="mw-redirect" title="Totipotent">totipotent</a>. In mammals, only the zygote and subsequent <a href="/wiki/Blastomere" title="Blastomere">blastomeres</a> are totipotent, while in plants, many differentiated cells can become totipotent with simple laboratory techniques. A cell that can differentiate into all cell types of the adult organism is known as <a href="/wiki/Pluripotent" class="mw-redirect" title="Pluripotent">pluripotent</a>. Such cells are called <a href="/wiki/Meristem" title="Meristem">meristematic cells</a> in higher plants and <a href="/wiki/Embryonic_stem_cell" title="Embryonic stem cell">embryonic stem cells</a> in animals, though some groups report the presence of adult pluripotent cells. Virally induced expression of four transcription factors <a href="/wiki/Oct4" class="mw-redirect" title="Oct4">Oct4</a>, <a href="/wiki/Sox2" class="mw-redirect" title="Sox2">Sox2</a>, <a href="/wiki/C-Myc" class="mw-redirect" title="C-Myc">c-Myc</a>, and <a href="/wiki/Klf4" class="mw-redirect" title="Klf4">Klf4</a> (<a href="/wiki/Yamanaka_factors" class="mw-redirect" title="Yamanaka factors">Yamanaka factors</a>) is sufficient to create pluripotent (iPS) cells from adult <a href="/wiki/Fibroblast" title="Fibroblast">fibroblasts</a>.<a href="#cite_note-5">[5]</a> A <a href="/wiki/Multipotent" class="mw-redirect" title="Multipotent">multipotent</a> cell is one that can differentiate into multiple different, but closely related cell types.<a href="#cite_note-Schoeler-6">[6]</a> <a href="/wiki/Cell_potency#Oligopotent" title="Cell potency">Oligopotent cells</a> are more restricted than multipotent, but can still differentiate into a few closely related cell types.<a href="#cite_note-Schoeler-6">[6]</a> Finally, <a href="/wiki/Unipotent" title="Unipotent">unipotent</a> cells can differentiate into only one cell type, but are capable of <a href="/wiki/Stem_cell_self-renewal" class="mw-redirect" title="Stem cell self-renewal">self-renewal</a>.<a href="#cite_note-Schoeler-6">[6]</a> In <a href="/wiki/Cytopathology" title="Cytopathology">cytopathology</a>, the level of cellular differentiation is used as a measure of <a href="/wiki/Cancer" title="Cancer">cancer</a> progression. "<a href="/wiki/Grading_(tumors)" title="Grading (tumors)">Grade</a>" is a marker of how differentiated a cell in a tumor is.<a href="#cite_note-7">[7]</a> <div id="toc" class="toc" role="navigation" aria-labelledby="mw-toc-heading"><input type="checkbox" role="button" id="toctogglecheckbox" class="toctogglecheckbox" style="display:none"><div class="toctitle" lang="en" dir="ltr"><h2 id="mw-toc-heading">Contents</h2><label class="toctogglelabel" for="toctogglecheckbox"></label></div> <ul> <li class="toclevel-1 tocsection-1"><a href="#Mammalian_cell_types">1 Mammalian cell types</a></li> <li class="toclevel-1 tocsection-2"><a href="#Dedifferentiation">2 Dedifferentiation</a></li> <li class="toclevel-1 tocsection-3"><a href="#Mechanisms">3 Mechanisms</a></li> <li class="toclevel-1 tocsection-4"><a href="#Epigenetic_control">4 Epigenetic control</a> <ul> <li class="toclevel-2 tocsection-5"><a href="#Importance_of_epigenetic_control">4.1 Importance of epigenetic control</a></li> <li class="toclevel-2 tocsection-6"><a href="#Mechanisms_of_epigenetic_regulation">4.2 Mechanisms of epigenetic regulation</a> <ul> <li class="toclevel-3 tocsection-7"><a href="#Pioneer_factors_(Oct4,_Sox2,_Nanog)">4.2.1 Pioneer factors (Oct4, Sox2, Nanog)</a></li> <li class="toclevel-3 tocsection-8"><a href="#Polycomb_repressive_complex_(PRC2)">4.2.2 Polycomb repressive complex (PRC2)</a></li> <li class="toclevel-3 tocsection-9"><a href="#Trithorax_group_proteins_(TrxG)">4.2.3 Trithorax group proteins (TrxG)</a></li> <li class="toclevel-3 tocsection-10"><a href="#DNA_methylation">4.2.4 DNA methylation</a></li> <li class="toclevel-3 tocsection-11"><a href="#Nucleosome_positioning">4.2.5 Nucleosome positioning</a> <ul> <li class="toclevel-4 tocsection-12"><a href="#Histone_acetylation_and_methylation">4.2.5.1 Histone acetylation and methylation</a></li> <li class="toclevel-4 tocsection-13"><a href="#In_stem_cells">4.2.5.2 In stem cells</a></li> </ul> </li> </ul> </li> <li class="toclevel-2 tocsection-14"><a href="#Role_of_signaling_in_epigenetic_control">4.3 Role of signaling in epigenetic control</a></li> <li class="toclevel-2 tocsection-15"><a href="#Effect_of_matrix_elasticity">4.4 Effect of matrix elasticity</a></li> </ul> </li> <li class="toclevel-1 tocsection-16"><a href="#Evolutionary_history">5 Evolutionary history</a></li> <li class="toclevel-1 tocsection-17"><a href="#See_also">6 See also</a></li> <li class="toclevel-1 tocsection-18"><a href="#References">7 References</a></li> </ul> </div> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(1)"><h2 id="Mammalian_cell_types">Mammalian cell types</h2> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=1" title="Edit section: Mammalian cell types" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-1 collapsible-block" id="mf-section-1"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/List_of_distinct_cell_types_in_the_adult_human_body" class="mw-redirect" title="List of distinct cell types in the adult human body">List of distinct cell types in the adult human body</a></div> Three basic categories of cells make up the mammalian body: <a href="/wiki/Germ_cell" title="Germ cell">germ cells</a>, <a href="/wiki/Somatic_cell" title="Somatic cell">somatic cells</a>, and <a href="/wiki/Stem_cell" title="Stem cell">stem cells</a>. Each of the approximately 37.2 trillion (3.72x1013) cells in an adult human has its own copy or copies of the <a href="/wiki/Genome" title="Genome">genome</a> except certain <a href="/wiki/Cell_types" class="mw-redirect" title="Cell types">cell types</a>, such as <a href="/wiki/Red_blood_cell" title="Red blood cell">red blood cells</a>, that lack nuclei in their fully differentiated state. Most cells are <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a>; they have two copies of each <a href="/wiki/Chromosome" title="Chromosome">chromosome</a>. Such cells, called somatic cells, make up most of the human body, such as skin and muscle cells. Cells differentiate to specialize for different functions.<a href="#cite_note-8">[8]</a> Germ line cells are any line of cells that give rise to <a href="/wiki/Gametes" class="mw-redirect" title="Gametes">gametes</a>—eggs and sperm—and thus are continuous through the generations. Stem cells, on the other hand, have the ability to divide for indefinite periods and to give rise to specialized cells. They are best described in the context of normal human development.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Development begins when a <a href="/wiki/Sperm" title="Sperm">sperm</a> fertilizes an <a href="/wiki/Egg_(biology)" class="mw-redirect" title="Egg (biology)">egg</a> and creates a single cell that has the potential to form an entire organism. In the first hours after fertilization, this cell divides into identical cells. In humans, approximately four days after fertilization and after several cycles of cell division, these cells begin to specialize, forming a hollow sphere of cells, called a <a href="/wiki/Blastocyst" title="Blastocyst">blastocyst</a>.<a href="#cite_note-9">[9]</a> The blastocyst has an outer layer of cells, and inside this hollow sphere, there is a cluster of cells called the <a href="/wiki/Inner_cell_mass" title="Inner cell mass">inner cell mass</a>. The cells of the inner cell mass go on to form virtually all of the tissues of the human body. Although the cells of the inner cell mass can form virtually every type of cell found in the human body, they cannot form an organism. These cells are referred to as <a href="/wiki/Pluripotent" class="mw-redirect" title="Pluripotent">pluripotent</a>.<a href="#cite_note-10">[10]</a> Pluripotent stem cells undergo further specialization into <a href="/wiki/Multipotent" class="mw-redirect" title="Multipotent">multipotent</a> <a href="/wiki/Progenitor_cell" title="Progenitor cell">progenitor cells</a> that then give rise to functional cells. Examples of stem and progenitor cells include:[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <ul><li><a href="/wiki/Radial_glial_cell" title="Radial glial cell">Radial glial cells</a> (embryonic neural stem cells) that give rise to excitatory neurons in the fetal brain through the process of <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a>.<a href="#cite_note-11">[11]</a><a href="#cite_note-12">[12]</a><a href="#cite_note-13">[13]</a></li> <li><a href="/wiki/Hematopoietic_stem_cells" class="mw-redirect" title="Hematopoietic stem cells">Hematopoietic stem cells</a> (adult stem cells) from the <a href="/wiki/Bone_marrow" title="Bone marrow">bone marrow</a> that give rise to red blood cells, <a href="/wiki/White_blood_cell" title="White blood cell">white blood cells</a>, and <a href="/wiki/Platelet" title="Platelet">platelets</a>.</li> <li><a href="/wiki/Mesenchymal_stem_cells" class="mw-redirect" title="Mesenchymal stem cells">Mesenchymal stem cells</a> (adult stem cells) from the bone marrow that give rise to stromal cells, fat cells, and types of bone cells</li> <li><a href="/wiki/Epithelia" class="mw-redirect" title="Epithelia">Epithelial</a> stem cells (progenitor cells) that give rise to the various types of skin cells</li> <li>Muscle <a href="/wiki/Satellite_cell" class="mw-redirect" title="Satellite cell">satellite cells</a> (progenitor cells) that contribute to differentiated <a href="/wiki/Muscle_tissue" class="mw-redirect" title="Muscle tissue">muscle tissue</a>.</li></ul> A pathway that is guided by the cell adhesion molecules consisting of four amino acids, <a href="/wiki/Arginine" title="Arginine">arginine</a>, <a href="/wiki/Glycine" title="Glycine">glycine</a>, <a href="/wiki/Asparagine" title="Asparagine">asparagine</a>, and <a href="/wiki/Serine" title="Serine">serine</a>, is created as the cellular blastomere <a href="/wiki/Gastrulation" title="Gastrulation">differentiates</a> from the single-layered <a href="/wiki/Blastula" class="mw-redirect" title="Blastula">blastula</a> to the three primary <a href="/wiki/Germ_layer" title="Germ layer">layers of germ cells</a> in mammals, namely the <a href="/wiki/Ectoderm" title="Ectoderm">ectoderm</a>, <a href="/wiki/Mesoderm" title="Mesoderm">mesoderm</a> and <a href="/wiki/Endoderm" title="Endoderm">endoderm</a> (listed from most distal (exterior) to proximal (interior)). The ectoderm ends up forming the skin and the nervous system, the mesoderm forms the bones and muscular tissue, and the endoderm forms the internal organ tissues. </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(2)"><h2 id="Dedifferentiation">Dedifferentiation</h2> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=2" title="Edit section: Dedifferentiation" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-2 collapsible-block" id="mf-section-2"> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Dedifferentiated_liposarcoma_-_intermed_mag.jpg" class="mw-file-description"><noscript><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Dedifferentiated_liposarcoma_-_intermed_mag.jpg/220px-Dedifferentiated_liposarcoma_-_intermed_mag.jpg" decoding="async" width="220" height="147" class="mw-file-element" data-file-width="4272" data-file-height="2848"></noscript><span class="lazy-image-placeholder" style="width: 220px;height: 147px;" data-src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Dedifferentiated_liposarcoma_-_intermed_mag.jpg/220px-Dedifferentiated_liposarcoma_-_intermed_mag.jpg" data-width="220" data-height="147" data-srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Dedifferentiated_liposarcoma_-_intermed_mag.jpg/330px-Dedifferentiated_liposarcoma_-_intermed_mag.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Dedifferentiated_liposarcoma_-_intermed_mag.jpg/440px-Dedifferentiated_liposarcoma_-_intermed_mag.jpg 2x" data-class="mw-file-element"> </a><figcaption><a href="/wiki/Micrograph" title="Micrograph">Micrograph</a> showing some dedifferentiation, (at left edge of image). + A differentiated component, showing <a href="/wiki/Lipoblast" title="Lipoblast">lipoblasts</a> and increased <a href="/wiki/Blood_vessels" class="mw-redirect" title="Blood vessels">vascularity</a>, (right edge of image). + Fully differentiated <a href="/wiki/Adipose_tissue" title="Adipose tissue">adipose tissue</a>, showing a few blood vessels, (center of image). (<a href="/wiki/Micrograph" title="Micrograph">Micrograph</a> of <a href="/wiki/Liposarcoma" title="Liposarcoma">liposarcoma</a> prepared with <a href="/wiki/H%26E_stain" title="H&E stain">H&E stain</a>).</figcaption></figure> <a href="/wiki/Dedifferentiation" title="Dedifferentiation">Dedifferentiation</a>, or integration, is a cellular process seen in the more <a href="/wiki/Basal_(phylogenetics)" title="Basal (phylogenetics)">basal</a> life forms in animals, such as <a href="/wiki/Worm" title="Worm">worms</a> and <a href="/wiki/Amphibian" title="Amphibian">amphibians</a> where a differentiated cell reverts to an earlier developmental stage—usually as part of a <a href="/wiki/Regeneration_(biology)" title="Regeneration (biology)">regenerative</a> process.<a href="#cite_note-dediff1-14">[14]</a><a href="#cite_note-dediff2-15">[15]</a> Dedifferentiation also occurs in plant cells.<a href="#cite_note-16">[16]</a> And, in <a href="/wiki/Cell_culture" title="Cell culture">cell culture</a> in the laboratory, cells can change shape or may lose specific properties such as protein expression—which processes are also termed dedifferentiation.<a href="#cite_note-17">[17]</a> Some hypothesize that dedifferentiation is an aberration that likely results in <a href="/wiki/Cancer" title="Cancer">cancers</a>,<a href="#cite_note-18">[18]</a> but others explain it as a natural part of the immune response that was lost to humans at some point of evolution. A newly discovered molecule dubbed <a href="/wiki/Reversine" title="Reversine">reversine</a>, a <a href="/wiki/Purine" title="Purine">purine</a> analog, has proven to induce dedifferentiation in <a href="/wiki/Myotube" class="mw-redirect" title="Myotube">myotubes</a>. These manifestly dedifferentiated cells—now performing essentially as stem cells—could then redifferentiate into <a href="/wiki/Osteoblast" title="Osteoblast">osteoblasts</a> and <a href="/wiki/Adipocyte" title="Adipocyte">adipocytes</a>.<a href="#cite_note-19">[19]</a> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Bischoff_SR_-_Nuclear_Reprogramming.pdf" class="mw-file-description"><noscript><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/56/Bischoff_SR_-_Nuclear_Reprogramming.pdf/page1-401px-Bischoff_SR_-_Nuclear_Reprogramming.pdf.jpg" decoding="async" width="401" height="310" class="mw-file-element" data-file-width="1650" data-file-height="1275"></noscript><span class="lazy-image-placeholder" style="width: 401px;height: 310px;" data-src="//upload.wikimedia.org/wikipedia/commons/thumb/5/56/Bischoff_SR_-_Nuclear_Reprogramming.pdf/page1-401px-Bischoff_SR_-_Nuclear_Reprogramming.pdf.jpg" data-width="401" data-height="310" data-srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/56/Bischoff_SR_-_Nuclear_Reprogramming.pdf/page1-602px-Bischoff_SR_-_Nuclear_Reprogramming.pdf.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/56/Bischoff_SR_-_Nuclear_Reprogramming.pdf/page1-802px-Bischoff_SR_-_Nuclear_Reprogramming.pdf.jpg 2x" data-class="mw-file-element"> </a><figcaption>Diagram exposing several methods used to revert adult somatic cells to <a href="/wiki/Totipotency" class="mw-redirect" title="Totipotency">totipotency</a> or <a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">pluripotency</a>.</figcaption></figure> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(3)"><h2 id="Mechanisms">Mechanisms</h2> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=3" title="Edit section: Mechanisms" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-3 collapsible-block" id="mf-section-3"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Embryonic_differentiation_waves" title="Embryonic differentiation waves">Embryonic differentiation waves</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cell_Differentiation.jpg" class="mw-file-description"><noscript><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/Cell_Differentiation.jpg/330px-Cell_Differentiation.jpg" decoding="async" width="330" height="174" class="mw-file-element" data-file-width="727" data-file-height="384"></noscript><span class="lazy-image-placeholder" style="width: 330px;height: 174px;" data-src="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/Cell_Differentiation.jpg/330px-Cell_Differentiation.jpg" data-width="330" data-height="174" data-srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/Cell_Differentiation.jpg/495px-Cell_Differentiation.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/64/Cell_Differentiation.jpg/660px-Cell_Differentiation.jpg 2x" data-class="mw-file-element"> </a><figcaption>Mechanisms of cellular differentiation</figcaption></figure> Each specialized cell type in an organism <a href="/wiki/Gene_expression" title="Gene expression">expresses</a> a <a href="/wiki/Subset" title="Subset">subset</a> of all the <a href="/wiki/Gene" title="Gene">genes</a> that constitute the genome of that <a href="/wiki/Species" title="Species">species</a>. Each cell type is defined by its particular pattern of <a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">regulated gene expression</a>. Cell differentiation is thus a transition of a cell from one cell type to another and it involves a switch from one pattern of gene expression to another. Cellular differentiation during development can be understood as the result of a <a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">gene regulatory network</a>. A regulatory gene and its cis-regulatory modules are nodes in a gene regulatory network; they receive input and create output elsewhere in the network.<a href="#cite_note-DeLeon-20">[20]</a> The <a href="/wiki/Systems_biology" title="Systems biology">systems biology</a> approach to developmental biology emphasizes the importance of investigating how developmental mechanisms interact to produce predictable patterns (<a href="/wiki/Morphogenesis" title="Morphogenesis">morphogenesis</a>). However, an alternative view has been proposed recently[<a href="/wiki/Wikipedia:Manual_of_Style/Dates_and_numbers#Chronological_items" title="Wikipedia:Manual of Style/Dates and numbers">when?</a>][<a href="/wiki/Wikipedia:Manual_of_Style/Words_to_watch#Unsupported_attributions" title="Wikipedia:Manual of Style/Words to watch">by whom?</a>]. Based on <a href="/wiki/Stochastic" title="Stochastic">stochastic</a> gene expression, cellular differentiation is the result of a Darwinian selective process occurring among cells. In this frame, protein and gene networks are the result of cellular processes and not their cause.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Signal_transduction_pathways.svg" class="mw-file-description"><noscript><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/240px-Signal_transduction_pathways.svg.png" decoding="async" width="240" height="176" class="mw-file-element" data-file-width="1858" data-file-height="1364"></noscript><span class="lazy-image-placeholder" style="width: 240px;height: 176px;" data-src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/240px-Signal_transduction_pathways.svg.png" data-width="240" data-height="176" data-srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/360px-Signal_transduction_pathways.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/480px-Signal_transduction_pathways.svg.png 2x" data-class="mw-file-element"> </a><figcaption>An overview of major signal transduction pathways.</figcaption></figure> While <a href="/wiki/Evolution" title="Evolution">evolutionarily</a> conserved molecular processes are involved in the cellular mechanisms underlying these switches, in animal species these are very different from the well-characterized <a href="/wiki/Operon" title="Operon">gene regulatory mechanisms</a> of <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, and even from those of the animals' closest <a href="/wiki/Holozoa" title="Holozoa">unicellular relatives</a>.<a href="#cite_note-Newman-21">[21]</a> Specifically, cell differentiation in animals is highly dependent on <a href="/wiki/Biomolecular_condensate" title="Biomolecular condensate">biomolecular condensates</a> of regulatory proteins and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a> DNA sequences. Cellular differentiation is often controlled by <a href="/wiki/Cell_signaling" title="Cell signaling">cell signaling</a>. Many of the signal molecules that convey information from cell to cell during the control of cellular differentiation are called <a href="/wiki/Growth_factor" title="Growth factor">growth factors</a>. Although the details of specific <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a> pathways vary, these pathways often share the following general steps. A ligand produced by one cell binds to a receptor in the extracellular region of another cell, inducing a conformational change in the receptor. The shape of the cytoplasmic domain of the receptor changes, and the receptor acquires enzymatic activity. The receptor then catalyzes reactions that phosphorylate other proteins, activating them. A cascade of phosphorylation reactions eventually activates a dormant transcription factor or cytoskeletal protein, thus contributing to the differentiation process in the target cell.<a href="#cite_note-Gilbert-22">[22]</a> Cells and tissues can vary in competence, their ability to respond to external signals.<a href="#cite_note-Rudel-23">[23]</a> Signal induction refers to cascades of signaling events, during which a cell or tissue signals to another cell or tissue to influence its developmental fate.<a href="#cite_note-Rudel-23">[23]</a> Yamamoto and Jeffery<a href="#cite_note-Yamamoto-24">[24]</a> investigated the role of the lens in eye formation in cave- and surface-dwelling fish, a striking example of induction.<a href="#cite_note-Rudel-23">[23]</a> Through reciprocal transplants, Yamamoto and Jeffery<a href="#cite_note-Yamamoto-24">[24]</a> found that the lens vesicle of surface fish can induce other parts of the eye to develop in cave- and surface-dwelling fish, while the lens vesicle of the cave-dwelling fish cannot.<a href="#cite_note-Rudel-23">[23]</a> Other important mechanisms fall under the category of <a href="/wiki/Asymmetric_cell_division" title="Asymmetric cell division">asymmetric cell divisions</a>, divisions that give rise to daughter cells with distinct developmental fates. Asymmetric cell divisions can occur because of asymmetrically expressed maternal cytoplasmic determinants or because of signaling.<a href="#cite_note-Rudel-23">[23]</a> In the former mechanism, distinct daughter cells are created during <a href="/wiki/Cytokinesis" title="Cytokinesis">cytokinesis</a> because of an uneven distribution of regulatory molecules in the parent cell; the distinct cytoplasm that each daughter cell inherits results in a distinct pattern of differentiation for each daughter cell. A well-studied example of pattern formation by asymmetric divisions is <a href="/wiki/Drosophila_embryogenesis#Anterior-posterior_axis_patterning_in_Drosophila" title="Drosophila embryogenesis">body axis patterning in Drosophila</a>. <a href="/wiki/RNA" title="RNA">RNA</a> molecules are an important type of intracellular differentiation control signal. The molecular and genetic basis of asymmetric cell divisions has also been studied in green algae of the genus <a href="/wiki/Volvox" title="Volvox">Volvox</a>, a model system for studying how unicellular organisms can evolve into multicellular organisms.<a href="#cite_note-Rudel-23">[23]</a> In Volvox carteri, the 16 cells in the anterior hemisphere of a 32-cell embryo divide asymmetrically, each producing one large and one small daughter cell. The size of the cell at the end of all cell divisions determines whether it becomes a specialized germ or somatic cell.<a href="#cite_note-Rudel-23">[23]</a><a href="#cite_note-Kirk-25">[25]</a> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(4)"><h2 id="Epigenetic_control">Epigenetic control</h2> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=4" title="Edit section: Epigenetic control" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-4 collapsible-block" id="mf-section-4"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Epigenetics_in_stem_cell_differentiation" class="mw-redirect" title="Epigenetics in stem cell differentiation">Epigenetics in stem cell differentiation</a></div> Since each cell, regardless of cell type, possesses the same genome, determination of cell type must occur at the level of gene expression. While the regulation of gene expression can occur through <a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-</a> and <a href="/wiki/Trans-regulatory_element" title="Trans-regulatory element">trans-regulatory elements</a> including a gene's <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoter</a> and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancers</a>, the problem arises as to how this expression pattern is maintained over numerous generations of <a href="/wiki/Cell_division" title="Cell division">cell division</a>.<a href="#cite_note-26">[26]</a> As it turns out, <a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">epigenetic</a> processes play a crucial role in regulating the decision to adopt a stem, progenitor, or mature <a href="/wiki/Cell_fate" class="mw-redirect" title="Cell fate">cell fate</a> This section will focus primarily on <a href="/wiki/Mammalian" class="mw-redirect" title="Mammalian">mammalian</a> <a href="/wiki/Stem_cells" class="mw-redirect" title="Stem cells">stem cells</a>. In systems biology and mathematical modeling of gene regulatory networks, cell-fate determination is predicted to exhibit certain dynamics, such as attractor-convergence (the attractor can be an equilibrium point, limit cycle or <a href="/wiki/Strange_attractor" class="mw-redirect" title="Strange attractor">strange attractor</a>) or oscillatory.<a href="#cite_note-27">[27]</a> <div class="mw-heading mw-heading3"><h3 id="Importance_of_epigenetic_control">Importance of epigenetic control</h3> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=5" title="Edit section: Importance of epigenetic control" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> The first question that can be asked is the extent and complexity of the role of epigenetic processes in the determination of cell fate. A clear answer to this question can be seen in the 2011 paper by Lister R, et al. <a href="#cite_note-Lister-28">[28]</a> on aberrant epigenomic programming in <a href="/wiki/Human" title="Human">human</a> <a href="/wiki/Induced_pluripotent_stem_cells" class="mw-redirect" title="Induced pluripotent stem cells">induced pluripotent stem cells</a>. As induced pluripotent stem cells (iPSCs) are thought to mimic <a href="/wiki/Embryonic_stem_cells" class="mw-redirect" title="Embryonic stem cells">embryonic stem cells</a> in their pluripotent properties, few epigenetic differences should exist between them. To test this prediction, the authors conducted whole-genome profiling of <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> patterns in several human embryonic stem cell (ESC), iPSC, and progenitor cell lines. Female <a href="/wiki/Adipose" class="mw-redirect" title="Adipose">adipose</a> cells, <a href="/wiki/Lung" title="Lung">lung</a> <a href="/wiki/Fibroblasts" class="mw-redirect" title="Fibroblasts">fibroblasts</a>, and foreskin fibroblasts were reprogrammed into induced pluripotent state with the <a href="/wiki/OCT4" class="mw-redirect" title="OCT4">OCT4</a>, <a href="/wiki/SOX2" title="SOX2">SOX2</a>, <a href="/wiki/KLF4" title="KLF4">KLF4</a>, and <a href="/wiki/MYC" title="MYC">MYC</a> genes. Patterns of DNA methylation in ESCs, iPSCs, somatic cells were compared. Lister R, et al. observed significant resemblance in methylation levels between embryonic and induced pluripotent cells. Around 80% of <a href="/wiki/CpG_site" title="CpG site">CG dinucleotides</a> in ESCs and iPSCs were methylated, the same was true of only 60% of CG dinucleotides in somatic cells. In addition, somatic cells possessed minimal levels of <a href="/wiki/Cytosine_methylation" class="mw-redirect" title="Cytosine methylation">cytosine methylation</a> in non-CG dinucleotides, while induced pluripotent cells possessed similar levels of methylation as embryonic stem cells, between 0.5 and 1.5%. Thus, consistent with their respective transcriptional activities,<a href="#cite_note-Lister-28">[28]</a> DNA methylation patterns, at least on the genomic level, are similar between ESCs and iPSCs. However, upon examining methylation patterns more closely, the authors discovered 1175 regions of differential CG dinucleotide methylation between at least one ES or iPS cell line. By comparing these regions of differential methylation with regions of cytosine methylation in the original somatic cells, 44-49% of differentially methylated regions reflected methylation patterns of the respective progenitor somatic cells, while 51-56% of these regions were dissimilar to both the progenitor and embryonic cell lines. <a href="/wiki/In_vitro" title="In vitro">In vitro</a>-induced differentiation of iPSC lines saw transmission of 88% and 46% of hyper and hypo-methylated differentially methylated regions, respectively. Two conclusions are readily apparent from this study. First, epigenetic processes are heavily involved in <a href="/wiki/Cell_fate_determination" title="Cell fate determination">cell fate determination</a>, as seen from the similar levels of cytosine methylation between induced pluripotent and embryonic stem cells, consistent with their respective patterns of <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a>. Second, the mechanisms of reprogramming (and by extension, differentiation) are very complex and cannot be easily duplicated, as seen by the significant number of differentially methylated regions between ES and iPS cell lines. Now that these two points have been established, we can examine some of the epigenetic mechanisms that are thought to regulate cellular differentiation. <div class="mw-heading mw-heading3"><h3 id="Mechanisms_of_epigenetic_regulation">Mechanisms of epigenetic regulation</h3> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=6" title="Edit section: Mechanisms of epigenetic regulation" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> <div class="mw-heading mw-heading4"><h4 id="Pioneer_factors_(Oct4,_Sox2,_Nanog)">Pioneer factors (Oct4, Sox2, Nanog)</h4> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=7" title="Edit section: Pioneer factors (Oct4, Sox2, Nanog)" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> Three transcription factors, OCT4, SOX2, and <a href="/wiki/Homeobox_protein_NANOG" title="Homeobox protein NANOG">NANOG</a> – the first two of which are used in induced pluripotent stem cell (iPSC) reprogramming, along with <a href="/wiki/Klf4" class="mw-redirect" title="Klf4">Klf4</a> and <a href="/wiki/C-Myc" class="mw-redirect" title="C-Myc">c-Myc</a> – are highly expressed in undifferentiated embryonic stem cells and are necessary for the maintenance of their <a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">pluripotency</a>.<a href="#cite_note-Christophersen-29">[29]</a> It is thought that they achieve this through alterations in <a href="/wiki/Chromatin" title="Chromatin">chromatin</a> structure, such as <a href="/wiki/Histone_modification" class="mw-redirect" title="Histone modification">histone modification</a> and DNA methylation, to restrict or permit the transcription of target genes. While highly expressed, their levels require a precise balance to maintain pluripotency, perturbation of which will promote differentiation towards different lineages based on how the gene expression levels change. Differential regulation of <a href="/wiki/Oct-4" title="Oct-4">Oct-4</a> and <a href="/wiki/SOX2" title="SOX2">SOX2</a> levels have been shown to precede germ layer fate selection.<a href="#cite_note-ReferenceA-30">[30]</a> Increased levels of Oct4 and decreased levels of Sox2 promote a <a href="/wiki/Gastrulation" title="Gastrulation">mesendodermal</a> fate, with Oct4 actively suppressing genes associated with a neural <a href="/wiki/Neurulation" title="Neurulation">ectodermal</a> fate. Similarly, increased levels of Sox2 and decreased levels of Oct4 promote differentiation towards a neural ectodermal fate, with Sox2 inhibiting differentiation towards a mesendodermal fate. Regardless of the lineage cells differentiate down, suppression of NANOG has been identified as a necessary prerequisite for differentiation.<a href="#cite_note-ReferenceA-30">[30]</a> <div class="mw-heading mw-heading4"><h4 id="Polycomb_repressive_complex_(PRC2)">Polycomb repressive complex (PRC2)</h4> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=8" title="Edit section: Polycomb repressive complex (PRC2)" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> In the realm of <a href="/wiki/Gene_silencing" title="Gene silencing">gene silencing</a>, <a href="/wiki/PRC2" title="PRC2">Polycomb repressive complex 2</a>, one of two classes of the <a href="/wiki/Polycomb-group_proteins" title="Polycomb-group proteins">Polycomb group</a> (PcG) family of proteins, catalyzes the di- and tri-methylation of histone H3 lysine 27 (H3K27me2/me3).<a href="#cite_note-Christophersen-29">[29]</a><a href="#cite_note-Jiang-31">[31]</a><a href="#cite_note-Guenther-32">[32]</a> By binding to the H3K27me2/3-tagged nucleosome, PRC1 (also a complex of PcG family proteins) catalyzes the mono-ubiquitinylation of histone H2A at lysine 119 (H2AK119Ub1), blocking <a href="/wiki/RNA_polymerase_II" title="RNA polymerase II">RNA polymerase II</a> activity and resulting in transcriptional suppression.<a href="#cite_note-Christophersen-29">[29]</a> PcG knockout ES cells do not differentiate efficiently into the three germ layers, and deletion of the PRC1 and PRC2 genes leads to increased expression of lineage-affiliated genes and unscheduled differentiation.<a href="#cite_note-Christophersen-29">[29]</a> Presumably, PcG complexes are responsible for transcriptionally repressing differentiation and development-promoting genes. <div class="mw-heading mw-heading4"><h4 id="Trithorax_group_proteins_(TrxG)">Trithorax group proteins (TrxG)</h4> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=9" title="Edit section: Trithorax group proteins (TrxG)" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> Alternately, upon receiving differentiation signals, PcG proteins are recruited to promoters of pluripotency transcription factors. PcG-deficient ES cells can begin differentiation but cannot maintain the differentiated phenotype.<a href="#cite_note-Christophersen-29">[29]</a> Simultaneously, differentiation and development-promoting genes are activated by Trithorax group (TrxG) chromatin regulators and lose their repression.<a href="#cite_note-Christophersen-29">[29]</a><a href="#cite_note-Guenther-32">[32]</a> TrxG proteins are recruited at regions of high transcriptional activity, where they catalyze the trimethylation of histone H3 lysine 4 (<a href="/wiki/H3K4me3" title="H3K4me3">H3K4me3</a>) and promote gene activation through histone acetylation.<a href="#cite_note-Guenther-32">[32]</a> PcG and TrxG complexes engage in direct competition and are thought to be functionally antagonistic, creating at differentiation and development-promoting loci what is termed a "bivalent domain" and rendering these genes sensitive to rapid induction or repression.<a href="#cite_note-Meissner-33">[33]</a> <div class="mw-heading mw-heading4"><h4 id="DNA_methylation">DNA methylation</h4> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=10" title="Edit section: DNA methylation" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> Regulation of gene expression is further achieved through DNA methylation, in which the <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">DNA methyltransferase</a>-mediated methylation of cytosine residues in CpG dinucleotides maintains heritable repression by controlling DNA accessibility.<a href="#cite_note-Meissner-33">[33]</a> The majority of CpG sites in embryonic stem cells are unmethylated and appear to be associated with H3K4me3-carrying nucleosomes.<a href="#cite_note-Christophersen-29">[29]</a> Upon differentiation, a small number of genes, including OCT4 and NANOG,<a href="#cite_note-Meissner-33">[33]</a> are methylated and their promoters repressed to prevent their further expression. Consistently, DNA methylation-deficient embryonic stem cells rapidly enter <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a> upon in vitro differentiation.<a href="#cite_note-Christophersen-29">[29]</a> <div class="mw-heading mw-heading4"><h4 id="Nucleosome_positioning">Nucleosome positioning</h4> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=11" title="Edit section: Nucleosome positioning" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> While the <a href="/wiki/Nucleic_acid_sequence" title="Nucleic acid sequence">DNA sequence</a> of most cells of an organism is the same, the binding patterns of transcription factors and the corresponding gene expression patterns are different. To a large extent, differences in transcription factor binding are determined by the chromatin accessibility of their binding sites through <a href="/wiki/Histone_modification" class="mw-redirect" title="Histone modification">histone modification</a> and/or <a href="/wiki/Pioneer_factor" title="Pioneer factor">pioneer factors</a>. In particular, it is important to know whether a <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a> is covering a given genomic binding site or not. This can be determined using a <a href="/wiki/Chromatin_immunoprecipitation" title="Chromatin immunoprecipitation">chromatin immunoprecipitation</a> assay.<a href="#cite_note-34">[34]</a> <div class="mw-heading mw-heading5"><h5 id="Histone_acetylation_and_methylation">Histone acetylation and methylation</h5> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=12" title="Edit section: Histone acetylation and methylation" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> DNA-nucleosome interactions are characterized by two states: either tightly bound by nucleosomes and transcriptionally inactive, called <a href="/wiki/Heterochromatin" title="Heterochromatin">heterochromatin</a>, or loosely bound and usually, but not always, transcriptionally active, called <a href="/wiki/Euchromatin" title="Euchromatin">euchromatin</a>. The epigenetic processes of histone methylation and acetylation, and their inverses demethylation and deacetylation primarily account for these changes. The effects of acetylation and deacetylation are more predictable. An acetyl group is either added to or removed from the positively charged Lysine residues in histones by enzymes called <a href="/wiki/Histone_acetyltransferase" title="Histone acetyltransferase">histone acetyltransferases</a> or <a href="/w/index.php?title=Histone_deactylase&action=edit&redlink=1" class="new" title="Histone deactylase (page does not exist)">histone deactylases</a>, respectively. The acetyl group prevents Lysine's association with the negatively charged DNA backbone. Methylation is not as straightforward, as neither methylation nor demethylation consistently correlate with either gene activation or repression. However, certain methylations have been repeatedly shown to either activate or repress genes. The trimethylation of lysine 4 on histone 3 (H3K4Me3) is associated with gene activation, whereas trimethylation of lysine 27 on histone 3 represses genes<a href="#cite_note-pmid12667454-35">[35]</a><a href="#cite_note-pmid12667453-36">[36]</a><a href="#cite_note-pmid15680324-37">[37]</a> <div class="mw-heading mw-heading5"><h5 id="In_stem_cells">In stem cells</h5> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=13" title="Edit section: In stem cells" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Stem_cell" title="Stem cell">Stem cell</a></div> During differentiation, stem cells change their gene expression profiles. Recent studies have implicated a role for nucleosome positioning and histone modifications during this process.<a href="#cite_note-Teif_et_al-38">[38]</a> There are two components of this process: turning off the expression of embryonic stem cell (ESC) genes, and the activation of cell fate genes. Lysine specific demethylase 1 (<a href="/wiki/LSD1" class="mw-redirect" title="LSD1">KDM1A</a>) is thought to prevent the use of <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a> regions of pluripotency genes, thereby inhibiting their transcription.<a href="#cite_note-ReferenceB-39">[39]</a> It interacts with <a href="/wiki/NuRD" class="mw-redirect" title="NuRD">Mi-2/NuRD complex</a> (nucleosome remodelling and histone deacetylase) complex,<a href="#cite_note-ReferenceB-39">[39]</a> giving an instance where methylation and acetylation are not discrete and mutually exclusive, but intertwined processes. <div class="mw-heading mw-heading3"><h3 id="Role_of_signaling_in_epigenetic_control">Role of signaling in epigenetic control</h3> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=14" title="Edit section: Role of signaling in epigenetic control" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> A final question to ask concerns the role of cell signaling in influencing the epigenetic processes governing differentiation. Such a role should exist, as it would be reasonable to think that extrinsic signaling can lead to epigenetic remodeling, just as it can lead to changes in gene expression through the activation or repression of different transcription factors. Little direct data is available concerning the specific signals that influence the <a href="/wiki/Epigenome" title="Epigenome">epigenome</a>, and the majority of current knowledge about the subject consists of speculations on plausible candidate regulators of epigenetic remodeling.<a href="#cite_note-Mohammad-40">[40]</a> We will first discuss several major candidates thought to be involved in the induction and maintenance of both embryonic stem cells and their differentiated progeny, and then turn to one example of specific signaling pathways in which more direct evidence exists for its role in epigenetic change. The first major candidate is <a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt signaling pathway</a>. The Wnt pathway is involved in all stages of differentiation, and the ligand Wnt3a can substitute for the overexpression of c-Myc in the generation of induced pluripotent stem cells.<a href="#cite_note-Mohammad-40">[40]</a> On the other hand, disruption of <a href="/wiki/Beta-catenin" class="mw-redirect" title="Beta-catenin">β-catenin</a>, a component of the Wnt signaling pathway, leads to decreased proliferation of neural progenitors. <a href="/wiki/Growth_factors" class="mw-redirect" title="Growth factors">Growth factors</a> comprise the second major set of candidates of epigenetic regulators of cellular differentiation. These morphogens are crucial for development, and include <a href="/wiki/Bone_morphogenetic_proteins" class="mw-redirect" title="Bone morphogenetic proteins">bone morphogenetic proteins</a>, <a href="/wiki/Transforming_growth_factors" class="mw-redirect" title="Transforming growth factors">transforming growth factors</a> (TGFs), and <a href="/wiki/Fibroblast_growth_factors" class="mw-redirect" title="Fibroblast growth factors">fibroblast growth factors</a> (FGFs). TGFs and FGFs have been shown to sustain expression of OCT4, SOX2, and NANOG by downstream signaling to <a href="/wiki/SMAD_(protein)" title="SMAD (protein)">Smad</a> proteins.<a href="#cite_note-Mohammad-40">[40]</a> Depletion of growth factors promotes the differentiation of ESCs, while genes with bivalent chromatin can become either more restrictive or permissive in their transcription.<a href="#cite_note-Mohammad-40">[40]</a> Several other signaling pathways are also considered to be primary candidates. Cytokine <a href="/wiki/Leukemia_inhibitory_factor" title="Leukemia inhibitory factor">leukemia inhibitory factors</a> are associated with the maintenance of mouse ESCs in an undifferentiated state. This is achieved through its activation of the Jak-STAT3 pathway, which has been shown to be necessary and sufficient towards maintaining mouse ESC pluripotency.<a href="#cite_note-Niwa-41">[41]</a> <a href="/wiki/Retinoic_acid" title="Retinoic acid">Retinoic acid</a> can induce differentiation of human and mouse ESCs,<a href="#cite_note-Mohammad-40">[40]</a> and <a href="/wiki/Notch_signaling" class="mw-redirect" title="Notch signaling">Notch signaling</a> is involved in the proliferation and self-renewal of stem cells. Finally, <a href="/wiki/Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">Sonic hedgehog</a>, in addition to its role as a morphogen, promotes embryonic stem cell differentiation and the self-renewal of somatic stem cells.<a href="#cite_note-Mohammad-40">[40]</a> The problem, of course, is that the candidacy of these signaling pathways was inferred primarily on the basis of their role in development and cellular differentiation. While epigenetic regulation is necessary for driving cellular differentiation, they are certainly not sufficient for this process. Direct modulation of gene expression through modification of transcription factors plays a key role that must be distinguished from heritable epigenetic changes that can persist even in the absence of the original environmental signals. Only a few examples of signaling pathways leading to epigenetic changes that alter cell fate currently exist, and we will focus on one of them. Expression of Shh (Sonic hedgehog) upregulates the production of <a href="/wiki/BMI1" title="BMI1">BMI1</a>, a component of the PcG complex that recognizes <a href="/wiki/H3K27me3" title="H3K27me3">H3K27me3</a>. This occurs in a Gli-dependent manner, as <a href="/wiki/Gli1" class="mw-redirect" title="Gli1">Gli1</a> and <a href="/wiki/Gli2" class="mw-redirect" title="Gli2">Gli2</a> are downstream effectors of the <a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a>. In culture, Bmi1 mediates the Hedgehog pathway's ability to promote human mammary stem cell self-renewal.<a href="#cite_note-Liu-42">[42]</a> In both humans and mice, researchers showed Bmi1 to be highly expressed in proliferating immature cerebellar granule cell precursors. When Bmi1 was knocked out in mice, impaired cerebellar development resulted, leading to significant reductions in postnatal brain mass along with abnormalities in motor control and behavior.<a href="#cite_note-Leung-43">[43]</a> A separate study showed a significant decrease in neural stem cell proliferation along with increased astrocyte proliferation in Bmi null mice.<a href="#cite_note-Zencak-44">[44]</a> An alternative model of cellular differentiation during embryogenesis is that positional information is based on mechanical signalling by the cytoskeleton using <a href="/wiki/Embryonic_differentiation_waves" title="Embryonic differentiation waves">Embryonic differentiation waves</a>. The mechanical signal is then epigenetically transduced via signal transduction systems (of which specific molecules such as Wnt are part) to result in differential gene expression. In summary, the role of signaling in the epigenetic control of cell fate in mammals is largely unknown, but distinct examples exist that indicate the likely existence of further such mechanisms. <div class="mw-heading mw-heading3"><h3 id="Effect_of_matrix_elasticity">Effect of matrix elasticity</h3> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=15" title="Edit section: Effect of matrix elasticity" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> In order to fulfill the purpose of regenerating a variety of tissues, adult stems are known to migrate from their niches, adhere to new extracellular matrices (ECM) and differentiate. The ductility of these microenvironments are unique to different tissue types. The ECM surrounding brain, muscle and bone tissues range from soft to stiff. The transduction of the stem cells into these cells types is not directed solely by chemokine cues and cell to cell signaling. The elasticity of the microenvironment can also affect the differentiation of mesenchymal stem cells (MSCs which originate in bone marrow.) When MSCs are placed on substrates of the same stiffness as brain, muscle and bone ECM, the MSCs take on properties of those respective cell types.<a href="#cite_note-:0-45">[45]</a> Matrix sensing requires the cell to pull against the matrix at focal adhesions, which triggers a cellular mechano-transducer to generate a signal to be informed what force is needed to deform the matrix. To determine the key players in matrix-elasticity-driven lineage specification in MSCs, different matrix microenvironments were mimicked. From these experiments, it was concluded that focal adhesions of the MSCs were the cellular mechano-transducer sensing the differences of the matrix elasticity. The non-muscle myosin IIa-c isoforms generates the forces in the cell that lead to signaling of early commitment markers. Nonmuscle myosin IIa generates the least force increasing to non-muscle myosin IIc. There are also factors in the cell that inhibit non-muscle myosin II, such as <a href="/wiki/Blebbistatin" title="Blebbistatin">blebbistatin</a>. This makes the cell effectively blind to the surrounding matrix.<a href="#cite_note-:0-45">[45]</a> Researchers have achieved some success in inducing stem cell-like properties in HEK 239 cells by providing a soft matrix without the use of diffusing factors.<a href="#cite_note-46">[46]</a> The stem-cell properties appear to be linked to tension in the cells' actin network. One identified mechanism for matrix-induced differentiation is tension-induced proteins, which remodel chromatin in response to mechanical stretch.<a href="#cite_note-47">[47]</a> The RhoA pathway is also implicated in this process.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(5)"><h2 id="Evolutionary_history">Evolutionary history</h2> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=16" title="Edit section: Evolutionary history" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-5 collapsible-block" id="mf-section-5"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Bangiomorpha" title="Bangiomorpha">Bangiomorpha</a></div> A billion-years-old, likely <a href="/wiki/Holozoa" title="Holozoa">holozoan</a>, <a href="/wiki/Protist" title="Protist">protist</a>, <a href="/wiki/Bicellum_brasieri" title="Bicellum brasieri">Bicellum brasieri</a> with two types of cells, shows that the evolution of differentiated <a href="/wiki/Multicellularity" class="mw-redirect" title="Multicellularity">multicellularity</a>, possibly but not necessarily of animal lineages, <a href="/wiki/History_of_life" title="History of life">occurred at least 1 billion years ago</a> and possibly mainly in <a href="/wiki/Freshwater_ecosystem" title="Freshwater ecosystem">freshwater lakes</a> rather than the ocean.<a href="#cite_note-48">[48]</a><a href="#cite_note-49">[49]</a><a href="#cite_note-50">[50]</a>[<a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify">clarification needed</a>] </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(6)"><h2 id="See_also">See also</h2> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=17" title="Edit section: See also" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-6 collapsible-block" id="mf-section-6"> <ul><li><a href="/wiki/Interbilayer_Forces_in_Membrane_Fusion" class="mw-redirect" title="Interbilayer Forces in Membrane Fusion">Interbilayer Forces in Membrane Fusion</a></li> <li><a href="/wiki/Fusion_mechanism" title="Fusion mechanism">Fusion mechanism</a></li> <li><a href="/wiki/Lipid_bilayer_fusion" title="Lipid bilayer fusion">Lipid bilayer fusion</a></li> <li><a href="/wiki/Cell-cell_fusogens" class="mw-redirect" title="Cell-cell fusogens">Cell-cell fusogens</a></li> <li><a href="/wiki/CAF-1" class="mw-redirect" title="CAF-1">CAF-1</a></li> <li><a href="/wiki/List_of_human_cell_types_derived_from_the_germ_layers" title="List of human cell types derived from the germ layers">List of human cell types derived from the germ layers</a></li></ul> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(7)"><h2 id="References">References</h2> <a role="button" href="/w/index.php?title=Cellular_differentiation&action=edit&section=18" title="Edit section: References" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-7 collapsible-block" id="mf-section-7"> <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-CME-1"><a href="#cite_ref-CME_1-0">^</a> <style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output 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<ul class="minerva-languages"><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AA%D9%85%D8%A7%D9%8A%D8%B2_%D8%AE%D9%84%D9%88%D9%8A" title="تمايز خلوي – Arabic" lang="ar" hreflang="ar" data-title="تمايز خلوي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/S%C3%A8-pau_hun-h%C3%B2a" title="Sè-pau hun-hòa – Minnan" lang="nan" hreflang="nan" data-title="Sè-pau hun-hòa" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9A%D0%BB%D0%B5%D1%82%D1%8A%D1%87%D0%BD%D0%BE_%D0%B4%D0%B8%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D0%B8%D1%80%D0%B0%D0%BD%D0%B5" title="Клетъчно диференциране – Bulgarian" lang="bg" hreflang="bg" data-title="Клетъчно диференциране" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/%C4%86elijska_diferencijacija" title="Ćelijska diferencijacija – Bosnian" lang="bs" hreflang="bs" data-title="Ćelijska diferencijacija" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Diferenciaci%C3%B3_cel%C2%B7lular" title="Diferenciació cel·lular – Catalan" lang="ca" hreflang="ca" data-title="Diferenciació cel·lular" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Bun%C4%9B%C4%8Dn%C3%A1_diferenciace" title="Buněčná diferenciace – Czech" lang="cs" hreflang="cs" data-title="Buněčná diferenciace" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Celledifferentiering" title="Celledifferentiering – Danish" lang="da" hreflang="da" data-title="Celledifferentiering" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Differenzierung_(Biologie)" title="Differenzierung (Biologie) – German" lang="de" hreflang="de" data-title="Differenzierung (Biologie)" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Diferentseerumine" title="Diferentseerumine – Estonian" lang="et" hreflang="et" data-title="Diferentseerumine" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9A%CF%85%CF%84%CF%84%CE%B1%CF%81%CE%B9%CE%BA%CE%AE_%CE%B4%CE%B9%CE%B1%CF%86%CE%BF%CF%81%CE%BF%CF%80%CE%BF%CE%AF%CE%B7%CF%83%CE%B7" title="Κυτταρική διαφοροποίηση – Greek" lang="el" hreflang="el" data-title="Κυτταρική διαφοροποίηση" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Diferenciaci%C3%B3n_celular" title="Diferenciación celular – Spanish" lang="es" hreflang="es" data-title="Diferenciación celular" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/%C4%88ela_diferenci%C4%9Do" title="Ĉela diferenciĝo – Esperanto" lang="eo" hreflang="eo" data-title="Ĉela diferenciĝo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Zelula_bereizte" title="Zelula bereizte – Basque" lang="eu" hreflang="eu" data-title="Zelula bereizte" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AA%D9%85%D8%A7%DB%8C%D8%B2_%D8%B3%D9%84%D9%88%D9%84%DB%8C" title="تمایز سلولی – Persian" lang="fa" hreflang="fa" data-title="تمایز سلولی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Diff%C3%A9renciation_cellulaire" title="Différenciation cellulaire – French" lang="fr" hreflang="fr" data-title="Différenciation cellulaire" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Diferenciaci%C3%B3n_celular" title="Diferenciación celular – Galician" lang="gl" hreflang="gl" data-title="Diferenciación celular" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%84%B8%ED%8F%AC_%EB%B6%84%ED%99%94" title="세포 분화 – Korean" lang="ko" hreflang="ko" data-title="세포 분화" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B2%D5%BB%D5%BB%D5%A1%D5%B5%D5%AB%D5%B6_%D5%BF%D5%A1%D6%80%D5%A2%D5%A5%D6%80%D5%A1%D5%AF%D5%B8%D6%82%D5%B4" title="Բջջային տարբերակում – Armenian" lang="hy" hreflang="hy" data-title="Բջջային տարբերակում" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Stani%C4%8Dna_diferencijacija" title="Stanična diferencijacija – Croatian" lang="hr" hreflang="hr" data-title="Stanična diferencijacija" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Diferensiasi_sel" title="Diferensiasi sel – Indonesian" lang="id" hreflang="id" data-title="Diferensiasi sel" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Differenziazione_cellulare" title="Differenziazione cellulare – Italian" lang="it" hreflang="it" data-title="Differenziazione cellulare" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%AA%D7%9E%D7%99%D7%99%D7%A0%D7%95%D7%AA_%D7%AA%D7%90%D7%99%D7%9D" title="התמיינות תאים – Hebrew" lang="he" hreflang="he" data-title="התמיינות תאים" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%A3%E1%83%AF%E1%83%A0%E1%83%94%E1%83%93%E1%83%94%E1%83%91%E1%83%98%E1%83%A1_%E1%83%93%E1%83%98%E1%83%A4%E1%83%94%E1%83%A0%E1%83%94%E1%83%9C%E1%83%AA%E1%83%98%E1%83%A0%E1%83%94%E1%83%91%E1%83%90" title="უჯრედების დიფერენცირება – Georgian" lang="ka" hreflang="ka" data-title="უჯრედების დიფერენცირება" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target">ქართული</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%96%D0%B0%D1%81%D1%83%D1%88%D0%B0%D0%BB%D0%B0%D1%80_%D0%B4%D0%B8%D1%84%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D0%B8%D1%8F%D1%81%D1%8B" title="Жасушалар дифференциясы – Kazakh" lang="kk" hreflang="kk" data-title="Жасушалар дифференциясы" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Pembezaan_sel" title="Pembezaan sel – Malay" lang="ms" hreflang="ms" data-title="Pembezaan sel" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Celdifferentiatie" title="Celdifferentiatie – Dutch" lang="nl" hreflang="nl" data-title="Celdifferentiatie" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%B4%B0%E8%83%9E%E5%88%86%E5%8C%96" title="細胞分化 – Japanese" lang="ja" hreflang="ja" data-title="細胞分化" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Celledifferensiering" title="Celledifferensiering – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Celledifferensiering" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Diferencia%C3%A7%C3%A3o_celular" title="Diferenciação celular – Portuguese" lang="pt" hreflang="pt" data-title="Diferenciação celular" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%94%D0%B8%D1%84%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D0%B8%D1%80%D0%BE%D0%B2%D0%BA%D0%B0_%D0%BA%D0%BB%D0%B5%D1%82%D0%BE%D0%BA" title="Дифференцировка клеток – Russian" lang="ru" hreflang="ru" data-title="Дифференцировка клеток" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Cell_differentiation" title="Cell differentiation – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cell differentiation" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Bunkov%C3%A1_diferenci%C3%A1cia" title="Bunková diferenciácia – Slovak" lang="sk" hreflang="sk" data-title="Bunková diferenciácia" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AC%DB%8C%D8%A7%D9%88%D8%A7%D8%B2%D8%A8%D9%88%D9%88%D9%86%DB%8C_%D8%AE%D8%A7%D9%86%DB%95%DB%8C" title="جیاوازبوونی خانەی – Central Kurdish" lang="ckb" hreflang="ckb" data-title="جیاوازبوونی خانەی" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%94%D0%B8%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D0%B8%D1%98%D0%B0%D1%86%D0%B8%D1%98%D0%B0_(%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%98%D0%B0)" title="Диференцијација (биологија) – Serbian" lang="sr" hreflang="sr" data-title="Диференцијација (биологија)" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Diferencijacija_(biologija)" title="Diferencijacija (biologija) – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Diferencijacija (biologija)" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Solujen_erilaistuminen" title="Solujen erilaistuminen – Finnish" lang="fi" hreflang="fi" data-title="Solujen erilaistuminen" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Celldifferentiering" title="Celldifferentiering – Swedish" lang="sv" hreflang="sv" data-title="Celldifferentiering" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Pagkakaiba-ibang_pangsihay" title="Pagkakaiba-ibang pangsihay – Tagalog" lang="tl" hreflang="tl" data-title="Pagkakaiba-ibang pangsihay" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target">Tagalog</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%89%E0%AE%AF%E0%AE%BF%E0%AE%B0%E0%AE%A3%E0%AF%81_%E0%AE%B5%E0%AF%87%E0%AE%B1%E0%AF%8D%E0%AE%B1%E0%AF%81%E0%AE%AE%E0%AF%88%E0%AE%AA%E0%AF%8D%E0%AE%AA%E0%AE%BE%E0%AE%9F%E0%AF%81" title="உயிரணு வேற்றுமைப்பாடு – Tamil" lang="ta" hreflang="ta" data-title="உயிரணு வேற்றுமைப்பாடு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%94%D0%B8%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D1%96%D0%B0%D1%86%D1%96%D1%8F_%D0%BA%D0%BB%D1%96%D1%82%D0%B8%D0%BD" title="Диференціація клітин – Ukrainian" lang="uk" hreflang="uk" data-title="Диференціація клітин" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%AE%D9%84%DB%8C%D8%A7%D8%AA%DB%8C_%D8%AA%D9%85%D8%A7%D9%8A%D9%8F%D8%B2" title="خلیاتی تمايُز – Urdu" lang="ur" hreflang="ur" data-title="خلیاتی تمايُز" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target">اردو</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Bi%E1%BB%87t_h%C3%B3a_t%E1%BA%BF_b%C3%A0o" title="Biệt hóa tế bào – Vietnamese" lang="vi" hreflang="vi" data-title="Biệt hóa tế bào" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target">Tiếng Việt</a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%B4%B0%E8%83%9E%E5%88%86%E5%8C%96" title="細胞分化 – Cantonese" lang="yue" hreflang="yue" data-title="細胞分化" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target">粵語</a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%BB%86%E8%83%9E%E5%88%86%E5%8C%96" title="细胞分化 – Chinese" lang="zh" hreflang="zh" data-title="细胞分化" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target">中文</a></li></ul> </section> </div> <div class="minerva-footer-logo"><img src="/static/images/mobile/copyright/wikipedia-wordmark-en.svg" alt="Wikipedia" width="120" height="18" style="width: 7.5em; height: 1.125em;"/> </div> <ul id="footer-info" class="footer-info hlist hlist-separated"> <li 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