Editing Apoptosis (section)

==Discovery and etymology==
{{Main|History of apoptosis research}}

German scientist [[Carl Vogt]] was first to describe the principle of apoptosis in 1842. In 1885, anatomist [[Walther Flemming]] delivered a more precise description of the process of programmed cell death. However, it was not until 1965 that the topic was resurrected. While studying tissues using electron microscopy, [[John Kerr (pathologist)|John Kerr]] at the University of Queensland was able to distinguish apoptosis from traumatic cell death.<ref>{{cite journal | vauthors = Kerr JF | title = A histochemical study of hypertrophy and ischaemic injury of rat liver with special reference to changes in lysosomes | journal = The Journal of Pathology and Bacteriology | volume = 90 | issue = 2 | pages = 419–435 | date = October 1965 | pmid = 5849603 | doi = 10.1002/path.1700900210 }}</ref> Following the publication of a paper describing the phenomenon, Kerr was invited to join [[Alastair Currie]], as well as [[Andrew Wyllie (pathologist)|Andrew Wyllie]], who was Currie's graduate student,<ref name="AHW 1972">{{cite web|url=http://www.a-star.edu.sg/astar/biomed/action/biomed_dvp_abstract.do?id=2901ddeb02dH |title=Prof Andrew H. Wyllie – Lecture Abstract |access-date=2007-03-30 |author =Agency for Science, Technology and Research |archive-url=https://web.archive.org/web/20071113101931/http://www.a-star.edu.sg/astar/biomed/action/biomed_dvp_abstract.do?id=2901ddeb02dH |archive-date=2007-11-13 |url-status=dead }}</ref> at the [[University of Aberdeen]]. In 1972, the trio published a seminal article in the ''[[British Journal of Cancer]]''.<ref name="Kerr1972">{{cite journal | vauthors = Kerr JF, Wyllie AH, Currie AR | title = Apoptosis: a basic biological phenomenon with wide-ranging implications in tissue kinetics | journal = British Journal of Cancer | volume = 26 | issue = 4 | pages = 239–257 | date = August 1972 | pmid = 4561027 | pmc = 2008650 | doi = 10.1038/bjc.1972.33 }}</ref> Kerr had initially used the term programmed cell necrosis, but in the article, the process of natural cell death was called ''apoptosis''. Kerr, Wyllie and Currie credited James Cormack, a professor of Greek language at University of Aberdeen, with suggesting the term apoptosis. Kerr received the [[Paul Ehrlich and Ludwig Darmstaedter Prize]] on March 14, 2000, for his description of apoptosis. He shared the prize with Boston biologist [[H. Robert Horvitz]].<ref>{{cite journal | vauthors = O'Rourke MG, Ellem KA | title = John Kerr and apoptosis | journal = The Medical Journal of Australia | volume = 173 | issue = 11–12 | pages = 616–617 | year = 2000 | pmid = 11379508 | doi = 10.5694/j.1326-5377.2000.tb139362.x | s2cid = 38265127 }}</ref>

For many years, neither "apoptosis" nor  "programmed cell death" was a highly cited term. Two discoveries brought cell death from obscurity to a major field of research: identification of the first component of the cell death control and effector mechanisms, and linkage of abnormalities in cell death to human disease, in particular cancer. This occurred in 1988 when it was shown that BCL2, the gene responsible for follicular lymphoma, encoded a protein that inhibited cell death.<ref name=bcl2>{{cite journal |vauthors=Vaux DL, Cory S, Adams JM |title=Bcl-2 gene promotes haemopoietic cell survival and cooperates with c-myc to immortalize pre-B cells |journal=Nature |volume=335 |issue=6189 |pages=440–2 |date=September 1988 |pmid=3262202 |doi=10.1038/335440a0 |bibcode=1988Natur.335..440V |s2cid=23593952 }}</ref>

The 2002 [[Nobel Prize in Physiology or Medicine|Nobel Prize in Medicine]] was awarded to [[Sydney Brenner]], [[H. Robert Horvitz]] and [[John Sulston]] for their work identifying genes that control apoptosis. The genes were identified by studies in the nematode ''[[Caenorhabditis elegans|C. elegans]]'' and homologues of these genes function in humans to regulate apoptosis.<ref>{{Cite web |title=The Nobel Prize in Physiology or Medicine 2002 |url=https://www.nobelprize.org/prizes/medicine/2002/summary/#content |access-date=2025-01-13 |website=NobelPrize.org |language=en-US}}</ref>

[[File:John Sulston.jpg|thumb|upright|[[John Sulston]] won the [[Nobel Prize in Medicine]] in 2002, for his pioneering research on apoptosis.]]

In Greek, apoptosis translates to the "falling off" of leaves from a tree.{{sfn|Alberts|p=1021}} Cormack, professor of Greek language, reintroduced the term for medical use as it had a medical meaning for the Greeks over two thousand years before. [[Hippocrates]] used the term to mean "the falling off of the bones". [[Galen]] extended its meaning to "the dropping of the scabs". Cormack was no doubt aware of this usage when he suggested the name. Debate continues over the correct pronunciation, with opinion divided between a pronunciation with the second ''p'' silent ({{IPAc-en|æ|p|ə|ˈ|t|oʊ|s|ᵻ|s}} {{respell|ap-ə|TOH|sis}}<ref name=ahdictionary>{{cite web |title=The American Heritage Dictionary entry: apoptosis |url=https://ahdictionary.com/word/search.html?q=apoptosis |website=ahdictionary.com |publisher=Houghton Mifflin Harcourt Publishing Company |access-date=26 July 2021 |date=2020 |archive-date=26 July 2021 |archive-url=https://web.archive.org/web/20210726211047/https://ahdictionary.com/word/search.html?q=apoptosis |url-status=live }}</ref><ref name="aboutapop">{{cite web|url=http://www.nih.gov/sigs/aig/Aboutapo.html|title=About apoptosis|access-date=2006-12-15|year=1999|author=Apoptosis Interest Group|archive-url= https://web.archive.org/web/20061228100402/http://www.nih.gov/sigs/aig/Aboutapo.html|archive-date=28 December 2006|url-status=dead}}</ref>) and the second ''p'' pronounced ({{IPAc-en|ei|p|ə|p|ˈ|t|oʊ|s|ᵻ|s|audio=LL-Q1860 (eng)-NaomiAmethyst-apoptosis.wav}}).<ref name=ahdictionary/><ref name="webster.com">{{cite web|url=http://www.webster.com/dictionary/apoptosis|title=Definition of apoptosis|website=www.webster.com|access-date=2007-08-11|archive-url= https://web.archive.org/web/20070703020311/http://webster.com/dictionary/apoptosis|archive-date=2007-07-03|url-status=dead}}</ref> In English, the ''p'' of the Greek ''-pt-'' [[consonant cluster]] is typically silent at the beginning of a word (e.g. [[Pterodactylus|pterodactyl]], [[Ptolemy]]), but articulated when used in combining forms preceded by a vowel, as in [[helicopter]] or the orders of insects: [[Fly|diptera]], [[lepidoptera]], etc.

In the original Kerr, Wyllie & Currie paper,<ref name=Kerr1972/> there is a footnote regarding the pronunciation:

<blockquote>We are most grateful to Professor James Cormack of the Department of Greek, University of Aberdeen, for suggesting this term. The word "apoptosis" ({{lang|grc|ἀπόπτωσις}}) is used in Greek to describe the "dropping off" or "falling off" of petals from flowers, or leaves from trees. To show the derivation clearly, we propose that the stress should be on the penultimate syllable, the second half of the word being pronounced like "ptosis" (with the "p" silent), which comes from the same root "to fall", and is already used to describe the drooping of the upper eyelid.</blockquote>