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//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Selection_shadow.svg/440px-Selection_shadow.svg.png 2x" data-file-width="440" data-file-height="600" /></a><figcaption><b>(a)</b> The survival rate of a population decreases as individuals age, but the reproduction rate remains constant. <b>(b)</b> The reproduction probability peaks early in life at sexual maturity and then steadily decreases as an individual ages, with the remaining share of the population decreasing with age as they enter the selection shadow.</figcaption></figure> <p>The <b>selection shadow</b> is a concept involved with the <a href="/wiki/Senescence#Evolutionary_theories" title="Senescence">evolutionary theories of aging</a> that states that <a href="/wiki/Selection_pressures" class="mw-redirect" title="Selection pressures">selection pressures</a> on an individual decrease as an individual ages and passes <a href="/wiki/Sexual_maturity" title="Sexual maturity">sexual maturity</a>, resulting in a "shadow" of time where <a href="/wiki/Selective_fitness" class="mw-redirect" title="Selective fitness">selective fitness</a> is not considered. Over generations, this results in maladaptive mutations that accumulate later in life due to aging being non-adaptive toward reproductive fitness.<sup id="cite_ref-Nature_1-0" class="reference"><a href="#cite_note-Nature-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The concept was first worked out by <a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a> and <a href="/wiki/Peter_Medawar" title="Peter Medawar">Peter Medawar</a> in the 1940s, with Medawar creating the first <a href="/wiki/Graphical_model" title="Graphical model">graphical model</a>.<sup id="cite_ref-Nature_1-1" class="reference"><a href="#cite_note-Nature-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Model">Model</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Selection_shadow&action=edit&section=1" title="Edit section: Model"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The model developed by Medawar states that due to the dangerous conditions and pressures from the environment, including predators and diseases, most individuals in the wild die not long after sexual maturity. Therefore, there is a low probability for individuals to survive to an advanced age and suffer the effects related to <a href="/wiki/Aging" class="mw-redirect" title="Aging">aging</a>. In conjunction with this, the effects of natural selection decrease as age increases, so that later individual performance is ignored by selection forces.<sup id="cite_ref-Nature_1-2" class="reference"><a href="#cite_note-Nature-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This results in beneficial <a href="/wiki/Mutations" class="mw-redirect" title="Mutations">mutations</a> not being selected for if they only have a positive result later in life, along with later in life deleterious mutations not being selected against. Due to the fitness of an individual not being affected once it is past its reproductive prime, later mutations and effects are considered to be in the "shadow" of selection.<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> </p><p>This concept would later be adapted into Medawar's 1952 <a href="/wiki/Evolution_of_ageing#Mutation_accumulation" title="Evolution of ageing">mutation accumulation hypothesis</a>, which was itself expanded upon by <a href="/wiki/George_C._Williams_(biologist)" class="mw-redirect" title="George C. Williams (biologist)">George C. Williams</a> in his 1957 <a href="/wiki/Antagonistic_pleiotropy_hypothesis" title="Antagonistic pleiotropy hypothesis">antagonistic pleiotropy hypothesis</a>.<sup id="cite_ref-Nature_1-3" class="reference"><a href="#cite_note-Nature-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>A classical requirement and <a href="/wiki/Constraint_(mathematics)" title="Constraint (mathematics)">constraint</a> of the model is that the number of individuals within a population that live to reach <a href="/wiki/Senescence" title="Senescence">senescence</a> must be small in number. If this is not true for a population, then the effects of old age will not be under a selection shadow and instead affect adaptation and evolution of the population as a whole. At the same time, however, this requirement has been challenged by increasing evidence of senescence being more common in wild populations than previously expected, especially among birds and mammals, while the effects of the selection shadow remain present.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Medawar's_Test_Tube_model"><span id="Medawar.27s_Test_Tube_model"></span>Medawar's Test Tube model</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Selection_shadow&action=edit&section=2" title="Edit section: Medawar's Test Tube model"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Peter_Brian_Medawar.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ca/Peter_Brian_Medawar.jpg/220px-Peter_Brian_Medawar.jpg" decoding="async" width="220" height="311" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/c/ca/Peter_Brian_Medawar.jpg 1.5x" data-file-width="280" data-file-height="396" /></a><figcaption>Peter Brian Medawar, the scientist/biologist behind the test tube theory. This theory was created to demonstrate how older test tubes are continuously replaced when they are broken, thus decreasing older test tube population numbers as time progresses. The test tubes were symbols for individual subjects in a species.</figcaption></figure> <p>Medawar developed a theoretical model to demonstrate his thought process which explained that most animals will die before aging will be the ultimate cause for death in that animal. This would be from environmental factors such as large storms, drought, and fires, and predation. Medawar wanted to demonstrate this possibility by using test tubes to get his point across. The test tubes would be used to represent a population of species.<sup id="cite_ref-:0_4-0" class="reference"><a href="#cite_note-:0-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> If one of these test tubes were to theoretically break, this would represent an individual animal dying. Randomly, test tubes would then be broken in the population to keep his model realistic. The broken test tubes would be replaced with a new one, which represents a new animal being born into the population.<sup id="cite_ref-:0_4-1" class="reference"><a href="#cite_note-:0-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Over time, the model showed that test tubes over a certain age would decline in the population as new test tubes were put in. The overall results in Medawar’s thought model demonstrated an exponential decline in the survivor curve which resulted in the population having a half life.<sup id="cite_ref-:0_4-2" class="reference"><a href="#cite_note-:0-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The amount of older animals, or test tubes in the population would then be harder to maintain and ultimately die. Medawar created this model to ultimately explain what would realistically happen in actual life.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Criticism">Criticism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Selection_shadow&action=edit&section=3" title="Edit section: Criticism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some scientists, however, have criticized the idea of aging being non-adaptive, instead adopting the theory of "<a href="/w/index.php?title=Death_by_Design_(theory)&action=edit&redlink=1" class="new" title="Death by Design (theory) (page does not exist)">death by design</a>". This theory follows the work of <a href="/wiki/August_Weismann" title="August Weismann">August Weismann</a>, which states that aging specifically evolved as an <a href="/wiki/Adaptation" title="Adaptation">adaptation</a>, and disagrees with Medawar's model as a perceived oversimplification of the impact older organisms have on evolution. It is also claimed that older organisms have a higher reproductive capacity due to being better fit in order to reach their age, rather than their capacity being equal as in Medawar's calculations.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Selection_shadow&action=edit&section=4" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-Nature-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Nature_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Nature_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Nature_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Nature_1-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><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 .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFFabianFlatt2011" class="citation web cs1">Fabian, Daniel; Flatt, Thomas (2011). <a rel="nofollow" class="external text" href="http://www.nature.com/scitable/knowledge/library/the-evolution-of-aging-23651151">"The Evolution of Aging"</a>. <i><a href="/wiki/Scitable" class="mw-redirect" title="Scitable">Scitable</a></i>. <a href="/wiki/Nature_Publishing_Group" class="mw-redirect" title="Nature Publishing Group">Nature Publishing Group</a><span class="reference-accessdate">. Retrieved <span class="nowrap">May 20,</span> 2014</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Scitable&rft.atitle=The+Evolution+of+Aging&rft.date=2011&rft.aulast=Fabian&rft.aufirst=Daniel&rft.au=Flatt%2C+Thomas&rft_id=http%3A%2F%2Fwww.nature.com%2Fscitable%2Fknowledge%2Flibrary%2Fthe-evolution-of-aging-23651151&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASelection+shadow" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFlattSchmidt2009" class="citation journal cs1">Flatt, Thomas; Schmidt, Paul S. (July 18, 2009). <a rel="nofollow" class="external text" href="http://i122server.vu-wien.ac.at/pop/Flatt_website/FlattSchmidt09.pdf">"Integrating evolutionary and molecular genetics of aging"</a> <span class="cs1-format">(PDF)</span>. <i>Biochimica et Biophysica Acta (BBA) - General Subjects</i>. <b>1790</b> (10). <a href="/wiki/Elsevier" title="Elsevier">Elsevier</a>: <span class="nowrap">951–</span>962. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.bbagen.2009.07.010">10.1016/j.bbagen.2009.07.010</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2972575">2972575</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19619612">19619612</a><span class="reference-accessdate">. Retrieved <span class="nowrap">May 20,</span> 2014</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biochimica+et+Biophysica+Acta+%28BBA%29+-+General+Subjects&rft.atitle=Integrating+evolutionary+and+molecular+genetics+of+aging&rft.volume=1790&rft.issue=10&rft.pages=%3Cspan+class%3D%22nowrap%22%3E951-%3C%2Fspan%3E962&rft.date=2009-07-18&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2972575%23id-name%3DPMC&rft_id=info%3Apmid%2F19619612&rft_id=info%3Adoi%2F10.1016%2Fj.bbagen.2009.07.010&rft.aulast=Flatt&rft.aufirst=Thomas&rft.au=Schmidt%2C+Paul+S.&rft_id=http%3A%2F%2Fi122server.vu-wien.ac.at%2Fpop%2FFlatt_website%2FFlattSchmidt09.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASelection+shadow" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTurbillRuf2010" class="citation journal cs1">Turbill, Christopher; Ruf, Thomas (August 6, 2010). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2917356">"Senescence Is More Important in the Natural Lives of Long- Than Short-Lived Mammals"</a>. <i><a href="/wiki/PLOS_One" title="PLOS One">PLOS One</a></i>. <b>5</b> (8). <a href="/wiki/Public_Library_of_Science" class="mw-redirect" title="Public Library of Science">Public Library of Science</a>: e12019. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2010PLoSO...512019T">2010PLoSO...512019T</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pone.0012019">10.1371/journal.pone.0012019</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2917356">2917356</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/20700508">20700508</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+One&rft.atitle=Senescence+Is+More+Important+in+the+Natural+Lives+of+Long-+Than+Short-Lived+Mammals&rft.volume=5&rft.issue=8&rft.pages=e12019&rft.date=2010-08-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2917356%23id-name%3DPMC&rft_id=info%3Apmid%2F20700508&rft_id=info%3Adoi%2F10.1371%2Fjournal.pone.0012019&rft_id=info%3Abibcode%2F2010PLoSO...512019T&rft.aulast=Turbill&rft.aufirst=Christopher&rft.au=Ruf%2C+Thomas&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2917356&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASelection+shadow" class="Z3988"></span></span> </li> <li id="cite_note-:0-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-:0_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:0_4-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:0_4-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.programmed-aging.org/theories/medawar_hypothesis.html">"Medawar's Hypothesis - Aging has a negligible effect on fitness"</a>. <i>www.programmed-aging.org</i><span class="reference-accessdate">. Retrieved <span class="nowrap">2020-04-13</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=www.programmed-aging.org&rft.atitle=Medawar%27s+Hypothesis+-+Aging+has+a+negligible+effect+on+fitness&rft_id=http%3A%2F%2Fwww.programmed-aging.org%2Ftheories%2Fmedawar_hypothesis.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASelection+shadow" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLachowiczMiękisz2009" class="citation book cs1">Lachowicz, Miroslaw; Miękisz, Jacek (April 20, 2009). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=m7n18NlBYSoC"><i>From Genetics to Mathematics</i></a>. <a href="/wiki/Singapore" title="Singapore">Singapore</a>: <a href="/wiki/World_Scientific" title="World Scientific">World Scientific</a>. p. 39. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-9812837257" title="Special:BookSources/978-9812837257"><bdi>978-9812837257</bdi></a><span class="reference-accessdate">. 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