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[[File:Centriole-en.svg|thumb|The [[centriole]], an [[organelle]] involved in [[cell division]], is structurally inherited.]]
'''Structural inheritance''' or '''cortical inheritance''' is the transmission of aan [[epigenetics|epigenetic]] trait in a living [[organism]] by a self-perpetuating spatial structures. This is in contrast to the transmission of digital information such as is found in [[DNA]] sequences, which accounts for the vast majority of known [[genetics|genetic]] variation.
 
Examples of structural inheritance include the propagation of [[prion]]s, the infectious proteins of diseases such as [[scrapie]] (in sheep and goats), [[bovine spongiform encephalopathy]] ('mad cow disease') and [[Creutzfeld-JakobCreutzfeldt–Jakob disease]] (although the protein-only hypothesis of prion transmission has been considered contentious until recently.) .<ref name="pmid15272271">{{cite journal |authorvauthors=Soto C, Castilla J | title = The controversial protein-only hypothesis of prion propagation | journal = Nat. Med. | volume = 10 Suppl | issue =Suppl 7| pages = S63–7 |yeardate=July 2004 |month=July |pmid = 15272271 | doi = 10.1038/nm1069 | s2cid = 8710254 }}</ref> Prions based on heritable protein structure also exist in [[yeast]] .<ref name="pmid7569955">{{cite journal |doi=10.1126/science.270.5233.93 |authorvauthors=Masison DC, Wickner RB | title = Prion-inducing ___domain of yeast Ure2p and protease resistance of Ure2p in prion-containing cells | journal = Science | volume = 270 | issue = 5233 | pages = 93–5 |yeardate=October 1995 |month=October |pmid = 7569955 | doi = 10.1126/science.270.5233.93 | url =http https://www.sciencemagzenodo.org/cgirecord/pmidlookup?view1231041 | bibcode =long&pmid 1995Sci...270...93M | s2cid =7569955 42262547 }}</ref><ref name="pmid8973157">{{cite journal |doi=10.1016/0168-9525(96)10045-7 |authorvauthors=Tuite MF, Lindquist SL | title = Maintenance and inheritance of yeast prions | journal = Trends Genet. | volume = 12 | issue = 11 | pages = 467–71 |yeardate=November 1996 |month=November |pmid = 8973157 |url doi =http://linkinghub 10.elsevier.com/retrieve/pii1016/0168-9525(96)10045-7 }}</ref><ref name="pmid11573346">{{cite journal |authorvauthors=Serio TR, Cashikar AG, Kowal AS, Sawicki GJ, Lindquist SL | title = Self-perpetuating changes in Sup35 protein conformation as a mechanism of heredity in yeast | journal = Biochem. Soc. Symp. | volume = 68| issue = 68 | pages = 35–43 | year = 2001 | pmid = 11573346 | doi = 10.1042/bss0680035| s2cid = 20173430 }}</ref>. Structural inheritance has also been seen in the orientation of [[cilium|cilia]] in protozoans such as ''[[Paramecium]]''<ref name="pmid14294056">{{cite journal |doi=10.1073/pnas.53.2.275 |authorvauthors=Beisson J, Sonneborn TM | title = Cytoplasmic inheritance of the organization of the cell cortex in ''Parameciumparamecium Aurelia''aurelia |journal=[[Proc. Natl. Acad. Sci. U.S.A.]] |volume = 53 | issue = 2| pages = 275–82 |yeardate=February 1965 |month=February |pmid = 14294056 | pmc = 219507 | doi = 10.1073/pnas.53.2.275 | bibcode = 1965PNAS...53..275B | doi-access = free }}</ref> and ''[[Tetrahymena]]'',<ref name="Nelsen89">{{cite journal |authorvauthors=Nelsen EM, Frankel J, Jenkins LM | title = Non-genic inheritance of cellular handedness | journal = Development | volume = 105 | issue = 3 | pages = 447–56 |yeardate=March 1989 |month doi =March 10.1242/dev.105.3.447 | pmid = 2612360 | url = http://dev.biologists.org/cgicontent/pmidlookup?view=long&pmid=2612360105/3/447.full.pdf }} </ref> and 'handedness' of the spiral of the cell in ''Tetrahymena'',<ref name=Nelsen89/> and shells of snails. Some [[organelle]]s also have structural inheritance, such as the [[centriole]], and the [[cell (biology)|cell]] itself (defined by the [[plasma membrane]]) may also be an example of structural inheritance. To emphasize the difference of the molecular mechanism of structural inheritance from the canonical [[Base pair|Watson-Crick base pairing]] mechanism of transmission of genetic information, the term 'epigenetic templating' was introduced.<ref name="pmid16809769">{{cite journal |vauthors=Viens A, Mechold U, Brouillard F, Gilbert C, Leclerc P, Ogryzko V | title = Analysis of human histone H2AZ deposition in vivo argues against its direct role in epigenetic templating mechanisms | journal = Mol. Cell. Biol. | volume = 26 | issue = 14 | pages = 5325–35 |date=July 2006 | pmid = 16809769 | pmc = 1592707 | doi = 10.1128/MCB.00584-06 }}</ref><ref name="pmid18419815">{{cite journal | author = Ogryzko VV | title = Erwin Schroedinger, Francis Crick and epigenetic stability | journal = Biol. Direct | volume = 3 | pages = 15 | year = 2008 | pmid = 18419815 | pmc = 2413215 | doi = 10.1186/1745-6150-3-15 | doi-access = free }}</ref>
 
==History==
Various additional examples of structural inheritance are presented in the recent book [[Origination of Organismal Form]].
Structural inheritance was discovered by [[Tracy Sonneborn]], and other researchers, during his study on [[protozoa]] in the late 1930s. Sonneborn demonstrated during his research on [[Paramecium]] that the structure of the cortex was not dependent on genes, or the liquid cytoplasm, but in the cortical structure of the surface of the ciliates. Preexisting cell surface structures provided a template that was passed on for many generations.<ref name="pmid16554410">{{cite journal | author = Preer JR | title = Sonneborn and the cytoplasm | journal = Genetics | volume = 172 | issue = 3 | pages = 1373–7 |date=March 2006 | doi = 10.1093/genetics/172.3.1373 | pmid = 16554410 | pmc = 1456306 }}</ref>
 
John R. Preer, Jr., following up on Sonneborn's work, says, "The arrangement of surface structures is inherited, but how is not known, Macronuclei pass on many of their characteristics to new macronuclei, by an unknown and mysterious mechanism."<ref name="pmid9071578">{{cite journal | author = Preer JR | title = Whatever happened to paramecium genetics? | journal = Genetics | volume = 145 | issue = 2 | pages = 217–25 |date=February 1997 | doi = 10.1093/genetics/145.2.217 | pmid = 9071578 | pmc = 1207789 }}</ref>
==External links==
*{{cite journal |doi=10.1073/pnas.212504699 |last1=Lindquist |first1=S. L. |title=Colloquium: Self-Perpetuating Structural States in Biology, Disease, and Genetics |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=99 |issue=Suppl 4 |pages=16377–506 |year=2002 |month=December |url=http://www.pnas.org/content/vol99/suppl_4/ }}
 
Other researchers have come to the conclusion that "the phenomena of cortical inheritance (and related nongenic, epigenetic processes) remind us that the fundamental reproductive unit of life is not a nucleic acid molecule, but the remarkably versatile, intact, living cell."{{cn|date=August 2022}}
==Notes and references==
{{reflist}}
 
The study of structural inheritance is part of the [[extended evolutionary synthesis]].<ref>[http://extendedevolutionarysynthesis.com/structural-inheritance-the-parent-as-a-developmental-template/ "Structural inheritance: The parent as a developmental template"]. Extended Evolutionary Synthesis.</ref>
[[Category:Epigenetics]][[Cortical Inheritance]]
 
==In popular culture==
{{genetics-stub}}
An article in ''[[Newsweek]]'' mentions research that shows that "Some water fleas sport a spiny helmet that deters predators; others, with identical DNA sequences, have bare heads. What differs between the two is not their genes but their mothers' experiences. If mom had a run-in with predators, her offspring have helmets, an effect one wag called "bite the mother, fight the daughter." If mom lived her life unthreatened, her offspring have no helmets. Same DNA, different traits. Somehow, the experience of the mother, not only her DNA sequences, has been transmitted to her offspring."<ref>{{cite web|author=Sharon BegleyJanuary 17, 2009 |url=http://www.newsweek.com/2009/01/16/the-sins-of-the-fathers-take-2.html |title=Begley: Was Darwin Wrong About Evolution? |publisher=Newsweek |date=2009-01-17 |access-date=2011-06-30}}</ref>
 
Various additional examples of structural inheritance are presented in the recent book ''[[Origination of Organismal Form]]''.
 
== References ==
{{reflist|2}}
 
== Further reading ==
{{refbegin}}
* {{cite journal |vauthors=Lindquist SL, Henikoff S | title = Self-perpetuating structural states in biology, disease, and genetics | journal = Proc. Natl. Acad. Sci. U.S.A. | volume = 99 Suppl 4 | issue = 90004| pages = 16377 |date=December 2002 | pmid = 12475994 | pmc = 139896 | doi = 10.1073/pnas.212504699 | bibcode = 2002PNAS...9916377L | doi-access = free }}
{{refend}}
 
[[Category:Epigenetics]][[Cortical Inheritance]]
[[Category:Extended evolutionary synthesis]]