[[File:Centriole-en.svg|thumb|The [[centriole]], an [[organelle]] involved in [[cell division]], is structurally inherited.]]
{{Merge from|Cortical Inheritance|date=January 2011}}
'''Structural inheritance''' or '''cortical inheritance''' is the transmission of an [[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 | author vauthors= Soto C, Castilla J | title = The controversial protein-only hypothesis of prion propagation | journal = Nat. Med. | volume = 10 Suppl | issue = 7| pages = S63–7 | year date=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 | author vauthors= 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 | year date=October 1995 | monthpmid = October7569955 | pmiddoi = 756995510.1126/science.270.5233.93 | doiurl = 10https://zenodo.1016org/0168-9525(96)10045-7record/1231041 | bibcode = 1995Sci...270...93M | s2cid = 42262547 }}</ref><ref name="pmid8973157">{{cite journal | author vauthors= Tuite MF, Lindquist SL | title = Maintenance and inheritance of yeast prions | journal = Trends Genet. | volume = 12 | issue = 11 | pages = 467–71 | year date=November 1996 | month = November | pmid = 8973157 | doi = 10.1016/0168-9525(96)10045-7 }}</ref><ref name="pmid11573346">{{cite journal | author vauthors= 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 | author vauthors= Beisson J, Sonneborn TM | title = Cytoplasmic inheritance of the organization of the cell cortex in paramecium aurelia | journal = [[Proc. Natl. Acad. Sci. U.S.A.]] | volume = 53 | issue = 2| pages = 275–82 | year date=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 | author vauthors= Nelsen EM, Frankel J, Jenkins LM | title = Non-genic inheritance of cellular handedness | journal = Development | volume = 105 | issue = 3 | pages = 447–56 | year date=March 1989 | monthdoi = March10.1242/dev.105.3.447 | pmid = 2612360 | doi = | url = http://dev.biologists.org/content/105/3/447.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 'Epigeneticepigenetic templating' was introduced.<ref name="pmid16809769">{{cite journal | author 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 | year date=July 2006 | month = July | 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 | issue = | pages = 15 | year = 2008 | pmid = 18419815 | pmc = 2413215 | doi = 10.1186/1745-6150-3-15 | doi-access = free }}</ref>
==History==
'''Cortical Inheritance''' or [[structuralStructural 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 | year date=March 2006 | monthdoi = March10.1093/genetics/172.3.1373 | pmid = 16554410 | pmc = 1456306 | doi = | url = | issn = }}</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 | year date=February 1997 | monthdoi = February10.1093/genetics/145.2.217 | pmid = 9071578 | pmc = 1207789 | doi = | url = | issn = }}</ref>
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."<ref>{{cite web|url=http://science.jrank.org/pages/48371/Cortical-Inheritance.html |title=Cortical Inheritance – Paramecium, Tetrahymena:, Teutophrys, Dileptus, Paramecium:, Pattern Formation: Ciliate Studies and Models – Cell, Cells, Structures, Prion, Genetic, and Information |publisher=Science.jrank.org cn|date=August |accessdate=2011-06-302022}}</ref>
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>
==In Popular Culture==
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=by 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 |accessdate=2011-06-30}}</ref> ▼
==In popular culture==
▲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= by 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 | accessdateaccess-date=2011-06-30}}</ref>
Various additional examples of structural inheritance are presented in the recent book ''[[Origination of Organismal Form]]''.
== Further reading ==
{{refbegin}}
* {{cite journal | author 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 | year date=December 2002 | month = December | pmid = 12475994 | pmc = 139896 | doi = 10.1073/pnas.212504699 | bibcode = 2002PNAS...9916377L | doi-access = free }}
{{refend}}
[[Category:Epigenetics]]
[[Category:Extended evolutionary synthesis]]
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