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<title>Clonal selection</title>

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

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</style><div role="note" class="hatnote navigation-not-searchable">For the use of clonal selection in general horticulture, see <a href="Clonal_(plant)" class="mw-redirect" title="Clonal (plant)">Clonal (plant)</a>. For the use of clonal selection in viticulture, see <a href="Propagation_of_grapevines" title="Propagation of grapevines">Propagation of grapevines</a>.</div>

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<p>In <a href="Immunology" title="Immunology">immunology</a>, <b>clonal selection theory</b> explains the functions of cells of the <a href="Immune_system" title="Immune system">immune system</a> (<a href="Lymphocytes" class="mw-redirect" title="Lymphocytes">lymphocytes</a>) in response to specific <a href="Antigens" class="mw-redirect" title="Antigens">antigens</a> invading the body. The concept was introduced by Australian doctor <a href="Frank_Macfarlane_Burnet" class="mw-redirect" title="Frank Macfarlane Burnet">Frank Macfarlane Burnet</a> in 1957, in an attempt to explain the great diversity of <a href="Antibodies" class="mw-redirect" title="Antibodies">antibodies</a> formed during initiation of the <a href="Immune_response" title="Immune response">immune response</a>.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The theory has become the widely accepted model for how the human <a href="Immune_system" title="Immune system">immune system</a> responds to infection and how certain types of <a href="B_cells" class="mw-redirect" title="B cells">B</a> and <a href="T_lymphocytes" class="mw-redirect" title="T lymphocytes">T lymphocytes</a> are selected for destruction of specific <a href="Antigens" class="mw-redirect" title="Antigens">antigens</a>.<sup id="cite_ref-Rajewsky1996_3-0" class="reference"><a href="#cite_note-Rajewsky1996-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>

</p><p>The theory states that in a pre-existing group of lymphocytes (both B and T cells), a specific antigen activates (i.e. selects) only its counter-specific cell, which then induces that particular cell to multiply, producing identical <a href="Clone_(B-cell_biology)" class="mw-redirect" title="Clone (B-cell biology)">clones</a> for antibody production. This activation occurs in secondary lymphoid organs such as the <a href="Spleen" title="Spleen">spleen</a> and the <a href="Lymph_node" title="Lymph node">lymph nodes</a>.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> In short, the theory is an explanation of the mechanism for the generation of diversity of antibody specificity.<sup id="cite_ref-Jordan_2007_72–79_5-0" class="reference"><a href="#cite_note-Jordan_2007_72–79-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The first experimental evidence came in 1958, when <a href="Gustav_Nossal" title="Gustav Nossal">Gustav Nossal</a> and <a href="Joshua_Lederberg" title="Joshua Lederberg">Joshua Lederberg</a> showed that one B cell always produces only one antibody.<sup id="cite_ref-nossal_6-0" class="reference"><a href="#cite_note-nossal-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The idea turned out to be the foundation of molecular immunology, especially in <a href="Adaptive_immunity" class="mw-redirect" title="Adaptive immunity">adaptive immunity</a>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>

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

<p>The clonal selection theory can be summarised with the following four tenets:

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<ul><li>Each lymphocyte bears a single type of receptor with a unique specificity (generated by <a href="V(D)J_recombination" title="V(D)J recombination">V(D)J recombination</a>).</li>

<li>Receptor occupation is required for cell activation.</li>

<li>The differentiated <a href="Effector_cell" title="Effector cell">effector cells</a> derived from an activated lymphocyte bear receptors of identical specificity as the parent cell.</li>

<li>Those lymphocytes bearing receptors for self molecules (i.e., <a href="Endogenous" class="mw-redirect" title="Endogenous">endogenous</a> antigens produced within the body) are destroyed at an early stage.</li></ul>

<div class="mw-heading mw-heading2"><h2 id="Early_work">Early work</h2></div>

<p>In 1900, <a href="Paul_Ehrlich" title="Paul Ehrlich">Paul Ehrlich</a> proposed the so-called "side chain theory" of antibody production. According to it, certain cells exhibit on their surface different "<a href="Side_chain" title="Side chain">side chains</a>" (i.e. membrane-bound antibodies) able to react with different antigens. When an antigen is present, it binds to a matching side chain. Then the cell stops producing all other side chains and starts intensive synthesis and secretion of the antigen-binding side chain as a soluble antibody. Though distinct from clonal selection, Ehrlich's idea was a selection theory far more accurate than the instructive theories that dominated immunology in the next decades.

</p><p>In 1955, Danish immunologist <a href="Niels_Jerne" class="mw-redirect" title="Niels Jerne">Niels Jerne</a> put forward a hypothesis that there is already a vast array of soluble antibodies in the <a href="Blood_serum" class="mw-redirect" title="Blood serum">serum</a> prior to any infection. The entrance of an antigen into the body results in the selection of only one type of antibody to match it. This supposedly occurred by certain cells <a href="Phagocytosis" title="Phagocytosis">phagocytosing</a> the immune complexes and somehow replicating the antibody structure to produce more of it.<sup id="cite_ref-Burnet1976_8-0" class="reference"><a href="#cite_note-Burnet1976-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup>

</p><p>In 1957, <a href="David_W._Talmage" class="mw-redirect" title="David W. Talmage">David W. Talmage</a> hypothesized that antigens bind to antibodies on the surface of antibody-producing cells and "only those cells are selected for multiplication whose synthesized product has affinity for the antigen". The key difference from Ehrlich's theory was that every cell was presumed to synthesize only one sort of antibody. After antigen binding the cell proliferates, forming clones with identical antibodies.

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<div class="mw-heading mw-heading3"><h3 id="Burnet's_clonal_selection_theory">Burnet's clonal selection theory</h3></div>

<p>Later in 1957, Australian immunologist <a href="Frank_Macfarlane_Burnet" class="mw-redirect" title="Frank Macfarlane Burnet">Frank Macfarlane Burnet</a> published a paper titled "A modification of Jerne's theory of antibody production using the concept of clonal selection" in the rather obscure <i>Australian Journal of Science</i>. In it Burnet expanded the ideas of Talmage and named the resulting theory the "clonal selection theory". He further formalised the theory in his 1959 book <i>The Clonal Selection Theory of Acquired Immunity</i>. He explained <a href="Immunological_memory" title="Immunological memory">immunological memory</a> as the cloning of two types of lymphocyte. One clone acts immediately to combat infection whilst the other is longer lasting, remaining in the immune system for a long time and causing immunity to that antigen. According to Burnet's hypothesis, among antibodies are molecules that can probably correspond with varying degrees of precision to all, or virtually all, the <a href="Antigenic_determinant" class="mw-redirect" title="Antigenic determinant">antigenic determinants</a> that occur in biological material other than those characteristic of the body itself. Each type of pattern is a specific product of a clone of lymphocytes and it is the essence of the hypothesis that each cell automatically has available on its surface representative reactive sites equivalent to those of the <a href="Globulin" title="Globulin">globulin</a> they produce. When an antigen enters the blood or tissue fluids it is assumed that it will attach to the surface of any lymphocyte carrying reactive sites that correspond to one of its antigenic determinants. Then the cell is activated and undergoes proliferation to produce a variety of descendants. In this way, preferential proliferation is initiated of all those clones whose reactive sites correspond to the antigenic determinants on the antigens present in the body. The descendants are capable of active liberation of soluble antibody and lymphocytes, the same functions as the parental forms.<sup id="cite_ref-Jordan_2007_72–79_5-1" class="reference"><a href="#cite_note-Jordan_2007_72–79-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup>

</p><p>In 1958, <a href="Gustav_Nossal" title="Gustav Nossal">Gustav Nossal</a> and <a href="Joshua_Lederberg" title="Joshua Lederberg">Joshua Lederberg</a> showed that one B cell always produces only one antibody, which was the first direct evidence supporting the clonal selection theory.<sup id="cite_ref-nossal_6-1" class="reference"><a href="#cite_note-nossal-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup>

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<div class="mw-heading mw-heading2"><h2 id="Theories_supported_by_clonal_selection">Theories supported by clonal selection</h2></div>

<p>Burnet and <a href="Peter_Medawar" title="Peter Medawar">Peter Medawar</a> worked together on understanding <a href="Immunological_tolerance" class="mw-redirect" title="Immunological tolerance">immunological tolerance</a>, a phenomenon also explained by clonal selection. This is the organism's ability to tolerate the introduction of cells prior to the development of an immune response as long as it occurs early in the organism's development. There are a vast number of lymphocytes occurring in the immune system, ranging from cells that tolerate self tissue to cells that do not. However, only cells tolerant of self tissue survive the embryonic stage.<span class="cleanup-needed-content" style="padding-left:0.1em; padding-right:0.1em; color:var(--color-subtle, #54595d); border:1px solid var(--border-color-subtle, #c8ccd1);">If non-self tissue is introduced, lymphocytes that develop are the ones that include the non-self tissues as self tissue.</span>

</p><p>In 1959, Burnet proposed that under certain circumstances, tissues could be successfully transplanted into foreign recipients. This work has led to a much greater understanding of the immune system and also great advances in tissue transplantation. Burnet and Medawar shared the <a href="Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> in 1960.

</p><p>In 1974, <a href="Niels_Kaj_Jerne" title="Niels Kaj Jerne">Niels Kaj Jerne</a> proposed that the immune system functions as a network that is regulated via interactions between the variable parts of lymphocytes and their secreted molecules. <a href="Immune_network_theory" title="Immune network theory">Immune network theory</a> is firmly based on the concept of clonal selection. Jerne won the Nobel Prize in Physiology or Medicine in 1984, largely for his contributions to immune network theory.

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<ul><li><a href="Adaptive_immune_system" title="Adaptive immune system">Adaptive immune system</a></li>

<li><a href="Clonal_selection_algorithm" title="Clonal selection algorithm">Clonal selection algorithm</a></li>

<li><a href="Universal_Darwinism" title="Universal Darwinism">Universal Darwinism</a></li></ul>

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

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</style><cite id="CITEREFBurnet1976" class="citation journal cs1">Burnet, FM (1976). <a rel="nofollow" class="external text" href="https://doi.org/10.3322%2Fcanjclin.26.2.119">"A modification of Jerne's theory of antibody production using the concept of clonal selection"</a>. <i>CA: A Cancer Journal for Clinicians</i>. <b>26</b> (2): <span class="nowrap">119–</span>21. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.3322%2Fcanjclin.26.2.119">10.3322/canjclin.26.2.119</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/816431">816431</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:40609269">40609269</a>.</cite></span>

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<li id="cite_note-Rajewsky1996-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-Rajewsky1996_3-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFRajewsky1996" class="citation journal cs1">Rajewsky, Klaus (1996). "Clonal selection and learning in the antibody system". <i>Nature</i>. <b>381</b> (6585): <span class="nowrap">751–</span>758. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1996Natur.381..751R">1996Natur.381..751R</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F381751a0">10.1038/381751a0</a>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0028-0836">0028-0836</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/8657279">8657279</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4279640">4279640</a>.</cite></span>

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<li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><cite id="CITEREFMurphy2012" class="citation book cs1">Murphy, Kenneth (2012). <i>Janeway's Immunobiology 8th Edition</i>. New York, NY: Garland Science. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>9780815342434</bdi>.</cite></span>

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<li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><cite id="CITEREFMedzhitov2013" class="citation journal cs1">Medzhitov, R. (2013). <a rel="nofollow" class="external text" href="https://doi.org/10.4049%2Fjimmunol.1302427">"Pattern Recognition Theory and the Launch of Modern Innate Immunity"</a>. <i>The Journal of Immunology</i>. <b>191</b> (9): <span class="nowrap">4473–</span>4474. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.4049%2Fjimmunol.1302427">10.4049/jimmunol.1302427</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24141853">24141853</a>.</cite></span>

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<li id="cite_note-Burnet1976-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-Burnet1976_8-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFBurnet1976" class="citation journal cs1">Burnet, F. M. (1976). <a rel="nofollow" class="external text" href="https://doi.org/10.3322%2Fcanjclin.26.2.119">"A Modification of Jerne's Theory of Antibody Production using the Concept of Clonal Selection"</a>. <i>CA: A Cancer Journal for Clinicians</i>. <b>26</b> (2): <span class="nowrap">119–</span>121. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.3322%2Fcanjclin.26.2.119">10.3322/canjclin.26.2.119</a></span>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0007-9235">0007-9235</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/816431">816431</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:40609269">40609269</a>.</cite></span>

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<li id="cite_note-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-9">^</a></b></span> <span class="reference-text"><cite id="CITEREFHodgkinHeath,_William_RBaxter,_Alan_G2007" class="citation journal cs1">Hodgkin, Philip D; Heath, William R; Baxter, Alan G (2007). "The clonal selection theory: 50 years since the revolution". <i>Nature Immunology</i>. <b>8</b> (10): <span class="nowrap">1019–</span>1026. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fni1007-1019">10.1038/ni1007-1019</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17878907">17878907</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:29935594">29935594</a>.</cite></span>

</li>

</ol></div></div>

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

<ul><li><cite id="CITEREFPodolsky1997" class="citation book cs1">Podolsky, Alfred I. Tauber; Scott H. (1997). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=ul2nIQjK2pwC"><i>The Generation of Diversity : Clonal Selection Theory and the Rise of Molecular Immunology</i></a> (1st paperback ed.). Cambridge, Massachusetts: Harvard Univ. Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>0-674-00182-6</bdi>.</cite><span class="cs1-maint citation-comment"><code class="cs1-code">{{cite book}}</code>: CS1 maint: multiple names: authors list (link)</span></li>

<li>"Biology in Context - The Spectrum of Life" Authors, Peter Aubusson, Eileen Kennedy.</li>

<li><cite id="CITEREFForsdyke_D.R.1995" class="citation journal cs1">Forsdyke D.R. (1995). <a rel="nofollow" class="external text" href="https://doi.org/10.1096%2Ffasebj.9.2.7781918">"The Origins of the Clonal Selection Theory of Immunity"</a>. <i>FASEB Journal</i>. <b>9</b> (2): <span class="nowrap">164–</span>66. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1096%2Ffasebj.9.2.7781918">10.1096/fasebj.9.2.7781918</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/7781918">7781918</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:38467403">38467403</a>.</cite></li></ul>

<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>

<ul><li><a rel="nofollow" class="external text" href="http://www.wehi.edu.au/education/wehitv/the_immune_system/">Animation of clonal selection</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110706121622/http://wehi.edu.au/education/wehitv/the_immune_system/">Archived</a> 6 July 2011 at the <a href="Wayback_Machine" title="Wayback Machine">Wayback Machine</a> from the Walter & Elisa Hall institute.</li></ul>

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</style><div id="Lymphocytic_adaptive_immune_system_and_complement123" style="font-size:114%;margin:0 4em"><a href="Lymphocyte" title="Lymphocyte">Lymphocytic</a> <a href="Adaptive_immune_system" title="Adaptive immune system">adaptive immune system</a> and <a href="Complement_system" title="Complement system">complement</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lymphoid</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Antigens</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

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

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

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

<li><a href="Antigenic_variation" title="Antigenic variation">Antigenic variation</a></li></ul></li>

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

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

<ul><li><a href="Linear_epitope" title="Linear epitope">Linear</a></li>

<li><a href="Conformational_epitope" title="Conformational epitope">Conformational</a></li></ul></li>

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

<ul><li><a href="Antigen_presentation" title="Antigen presentation">Antigen presentation</a>/<a href="Antigen-presenting_cell" title="Antigen-presenting cell">professional APCs</a>: <a href="Dendritic_cell" title="Dendritic cell">Dendritic cell</a></li>

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

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

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Antibodies</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="Antibody" title="Antibody">Antibody</a>

<ul><li><a href="Monoclonal_antibody" title="Monoclonal antibody">Monoclonal antibodies</a></li>

<li><a href="Polyclonal_antibodies" title="Polyclonal antibodies">Polyclonal antibodies</a></li>

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

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

<li><a href="Polyclonal_B_cell_response" title="Polyclonal B cell response">Polyclonal B cell response</a></li>

<li><a href="Allotype_(immunology)" title="Allotype (immunology)">Allotype</a></li>

<li><a href="Isotype_(immunology)" title="Isotype (immunology)">Isotype</a></li>

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

<ul><li><a href="Immune_complex" title="Immune complex">Immune complex</a></li>

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

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

<ul><li>Action: <a href="Immunity_(medical)" class="mw-redirect" title="Immunity (medical)">Immunity</a></li>

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

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

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

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

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

<li><a href="Cross-reactivity" title="Cross-reactivity">Cross-reactivity</a></li>

<li><a href="Co-stimulation" title="Co-stimulation">Co-stimulation</a></li></ul>

<ul><li>Inaction: <a href="Immune_tolerance" title="Immune tolerance">Tolerance</a>

<ul><li><a href="Central_tolerance" title="Central tolerance">Central</a></li>

<li><a href="Peripheral_tolerance" title="Peripheral tolerance">Peripheral</a></li>

<li><a href="Clonal_anergy" title="Clonal anergy">Clonal anergy</a></li>

<li><a href="Clonal_deletion" title="Clonal deletion">Clonal deletion</a></li>

<li><a href="T-cell_depletion" title="T-cell depletion">T-cell depletion</a></li>

<li><a href="Immune_tolerance_in_pregnancy" title="Immune tolerance in pregnancy">Tolerance in pregnancy</a></li></ul></li>

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

<li><a href="Immune_privilege" title="Immune privilege">Immune privilege</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Immunogenetics" title="Immunogenetics">Immunogenetics</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="Affinity_maturation" title="Affinity maturation">Affinity maturation</a>

<ul><li><a href="Somatic_hypermutation" title="Somatic hypermutation">Somatic hypermutation</a></li>

</ul></li>

<li><a href="V(D)J_recombination" title="V(D)J recombination">V(D)J recombination</a></li>

<li><a href="Junctional_diversity" title="Junctional diversity">Junctional diversity</a></li>

<li><a href="Immunoglobulin_class_switching" title="Immunoglobulin class switching">Immunoglobulin class switching</a></li>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Lymphocyte" title="Lymphocyte">Lymphocytes</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="Cell-mediated_immunity" title="Cell-mediated immunity">Cellular</a>

<li><a href="Humoral_immunity" title="Humoral immunity">Humoral</a>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Substances</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="Cytokine" title="Cytokine">Cytokines</a></li>

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

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

</div></td></tr></tbody></table></div></div><div><div class="zim-footer">

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