Editing Mitochondrion (section)

==History==
The first observations of intracellular structures that probably represented mitochondria were published in 1857, by the physiologist [[Albert von Kölliker|Albert von Kolliker]].<ref>{{cite journal |vauthors=Kölliker A |title=Einige Bemerkungen über die Endigungen der Hautnerven und den Bau der Muskeln |journal=Zeitschrift für wissenschaftliche Zoologie |date=1857 |volume=8 |pages=311–325 |url=https://www.biodiversitylibrary.org/item/49190#page/319/mode/1up |trans-title=Some remarks about the terminations of the cutaneous nerves and the structure of muscles |language=German |access-date=June 22, 2023 |archive-date=June 22, 2023 |archive-url=https://web.archive.org/web/20230622033955/https://www.biodiversitylibrary.org/item/49190#page/319/mode/1up |url-status=live }}  On p. 316, Kölliker described mitochondria which he observed in fresh frog muscles:  ''" ... sehr blasse rundliche Körnchen, welche in langen linienförmigen Zügen [...] wenn man einmal auf dieselben aufmerksam geworden ist."''  ( ... [they are] very faint round granules, which are embedded in the [muscle's] contractile substance in long linear trains.  These granules are located in the whole thickness of the muscle fiber, on the surface as in the interior, and [they] are so numerous that they appear as a not unimportant element of the muscle fibers, once one has become alert to them.)  Kölliker said (p. 321) that he had found mitochondria in the muscles of other animals.  In Figure 3 of  [https://www.biodiversitylibrary.org/item/49190#page/564/mode/1up Table XIV], Kölliker depicted mitochondria in frog muscles.</ref><ref name="Ernster-1981">{{cite journal | vauthors = Ernster L, Schatz G | title = Mitochondria: a historical review | journal = The Journal of Cell Biology | volume = 91 | issue = 3 Pt 2 | pages = 227s–255s | date = December 1981 | pmid = 7033239 | pmc = 2112799 | doi = 10.1083/jcb.91.3.227s }}</ref> [[Richard Altmann]], in 1890, established them as cell organelles and called them "bioblasts".<ref name="Ernster-1981"/><ref>{{cite book |vauthors=Altmann R |title=Die Elementarorganismen und ihre Beziehungen zu den Zellen |trans-title=Elementary Organisms and Their Relations to Cells |date=1890 |publisher=Veit & Co. |location=Leipzig, Germany |page=125 |url=https://www.deutschestextarchiv.de/book/view/altmann_elementarorganismen_1890?p=141 |language=German |access-date=June 23, 2023 |archive-date=June 23, 2023 |archive-url=https://web.archive.org/web/20230623002552/https://www.deutschestextarchiv.de/book/view/altmann_elementarorganismen_1890?p=141 |url-status=live }}  From p. 125:  ''"Da auch sonst mancherlei Umstände dafür sprechen, dass Mikroorganismen und Granula einander gleichwerthig sind und Elementarorganismen vorstellen, welche sich überall finden, wo lebendige Kräfte ausgelöst werden, so wollen wir sie mit dem gemeinschaftlichen Namen der Bioblasten bezeichnen."'' (Since otherwise some circumstances indicate that microorganisms and granula are equivalent to each other and suggest elementary organisms, which are to be found wherever living forces are unleashed, we will designate them with the collective name of "bioblasts".)</ref> In 1898, Carl Benda coined the term "mitochondria" from the [[Greek language|Greek]] {{lang|el|μίτος}}, {{transliteration|el|mitos}}, "thread", and {{lang|el|χονδρίον}}, {{transliteration|el|chondrion}}, "granule".<ref name="OnlineEtymDict">{{cite encyclopedia|title=mitochondria|url=http://www.etymonline.com/index.php?term=mitochondria&allowed_in_frame=0|dictionary=[[Online Etymology Dictionary]]|access-date=May 23, 2013|archive-date=March 4, 2016|archive-url=https://web.archive.org/web/20160304105458/http://www.etymonline.com/index.php?term=mitochondria&allowed_in_frame=0|url-status=live}}</ref><ref name="Ernster-1981"/><ref>{{cite journal | vauthors = Benda C | date = 1898 | title = Ueber die Spermatogenese der Vertebraten und höherer Evertebraten. II. Theil: Die Histiogenese der Spermien. | trans-title = On spermatogenesis in vertebrates and higher invertebrates. Part II: The histogenesis of sperm. | journal = Archiv für Physiologie | volume = 1898 | pages = 393–398 | url = https://www.biodiversitylibrary.org/item/109725#page/403/mode/1up | language = de | access-date = January 14, 2018 | archive-date = February 24, 2019 | archive-url = https://web.archive.org/web/20190224231523/https://www.biodiversitylibrary.org/item/109725#page/403/mode/1up | url-status = live }}  From p. 397:  After Brenda states that ''" ... ich bereits in vielen Zellarten aller möglichen Thierclassen gefunden habe, ... "'' ( ... I have already found [them (mitochondria)] in many types of cells of all possible classes of animals, ... ), he suggests:  ''"Ich möchte vorläufig vorschlagen, ihnen als Mitochondria eine besondere Stellung vorzubehalten, die ich in weiteren Arbeiten begründen werde."''  (I would like to suggest provisionally reserving for them, as "mitochondria", a special status which I will justify in further work.)
* {{cite journal |vauthors=Benda C |title=Weitere Mitteilungen über die Mitochondria |journal=Archiv für Physiologie: Verhandlungen der Berliner Physiologischen Gesellschaft |date=1899 |volume=1899 |pages=376–383 |url=https://www.biodiversitylibrary.org/item/109704#page/388/mode/1up |trans-title=Further reports on mitochondria |language=German |access-date=June 23, 2023 |archive-date=June 23, 2023 |archive-url=https://web.archive.org/web/20230623035548/https://www.biodiversitylibrary.org/item/109704#page/388/mode/1up |url-status=live }}</ref> [[Leonor Michaelis]] discovered that [[Janus green]] can be used as a [[supravital stain]] for mitochondria in 1900.<ref>{{cite journal |vauthors=Michaelis L |title=Die vitale Farbung, eine Darstellungsmethode der Zellgranula |journal=Archiv für Mikroskopische Anatomie und Entwicklungsgeschichte |trans-journal=Archive for Microscopic Anatomy and Ontogenesis |date=1900 |volume=55 |pages=558–575 |doi=10.1007/BF02977747 }}</ref> In 1904, [[Friedrich Meves]] made the first recorded observation of mitochondria in plants in cells of the white waterlily, ''[[Nymphaea alba]],''<ref name="Ernster-1981"/><ref>Ernster's citation {{cite journal |vauthors=Meves F |date=May 1908 |title=Die Chondriosomen als Träger erblicher Anlagen. Cytologische Studien am Hühnerembryo |journal=Archiv für Mikroskopische Anatomie |volume=72 |issue=1 |pages=816–867 |doi=10.1007/BF02982402 }} is wrong, correct citation is {{cite journal | vauthors = Meves F |year= 1904 |title=Über das Vorkommen von Mitochondrien bezw. Chondromiten in Pflanzenzellen |journal=Ber. Dtsch. Bot. Ges. |volume=22 |pages=284–286|doi= 10.1111/j.1438-8677.1904.tb05237.x }}, cited in Meves' 1908 paper and in {{cite journal |vauthors=Schmidt EW |year=1913 |title=Pflanzliche Mitochondrien |journal=Progressus Rei Botanicae |volume=4 |pages=164–183 |url=https://archive.org/stream/progressusreibot04lots/progressusreibot04lots_djvu.txt |access-date=September 21, 2012 }}, with confirmation of Nymphaea alba<!-- I was unable to access the full article, so I cannot confirm if "Nymphea" is Nymphaea. --></ref> and in 1908, along with [[Claudius Regaud]], suggested that they contain proteins and lipids. Benjamin F. Kingsbury, in 1912, first related them with cell respiration, but almost exclusively based on morphological observations.<ref>{{cite journal |vauthors=Kingsbury BF |title=Cytoplasmic fixation |journal=The Anatomical Record |date=1912 |volume=6 |issue=2 |pages=39–52 |doi=10.1002/ar.1090060202 }} From p. 47: " ... the mitochondria are the structural expression thereof [i.e., of the chemical reducing processes in the cytoplasm], ... "</ref><ref name="Ernster-1981"/> In 1913, [[Otto Heinrich Warburg]] linked respiration to particles which he had obtained from extracts of guinea-pig liver and which he called "grana".<ref>{{cite journal |vauthors=Warburg O |title=Über sauerstoffatmende Körnchen aus Leberzellen und über Sauerstoffatmung in Berkefeld-Filtraten wässriger Leberextrake |journal=Pflügers Archiv für die gesamte Physiologie des Menschen und der Tiere (Pfluger's Archive for All Physiology of Humans and Animals) |date=1913 |volume=154 |pages=599–617 |url=https://www.biodiversitylibrary.org/item/108058#page/613/mode/1up |trans-title=On respiring granules from liver cells and on respiration in Berkefeld filtrates of aqueous liver extracts |language=German |access-date=June 23, 2023 |archive-date=June 23, 2023 |archive-url=https://web.archive.org/web/20230623190650/https://www.biodiversitylibrary.org/item/108058#page/613/mode/1up |url-status=live }}</ref> Warburg and [[Heinrich Otto Wieland]], who had also postulated a similar particle mechanism, disagreed on the chemical nature of the respiration. It was not until 1925, when [[David Keilin]] discovered [[cytochromes]], that the [[respiratory chain]] was described.<ref name="Ernster-1981"/>

In 1939, experiments using minced muscle cells demonstrated that cellular respiration using one [[Oxygen|oxygen molecule]] can form four [[adenosine triphosphate]] (ATP) molecules, and in 1941, the concept of the phosphate bonds of ATP being a form of energy in cellular metabolism was developed by [[Fritz Albert Lipmann]]. In the following years, the mechanism behind cellular respiration was further elaborated, although its link to the mitochondria was not known.<ref name="Ernster-1981"/> The introduction of [[Cell fractionation|tissue fractionation]] by [[Albert Claude]] allowed mitochondria to be isolated from other cell fractions and biochemical analysis to be conducted on them alone. In 1946, he concluded that [[cytochrome oxidase]] and other enzymes responsible for the respiratory chain were isolated to the mitochondria. [[Eugene P. Kennedy|Eugene Kennedy]] and [[Albert L. Lehninger|Albert Lehninger]] discovered in 1948 that mitochondria are the site of [[oxidative phosphorylation]] in eukaryotes. Over time, the fractionation method was further developed, improving the quality of the mitochondria isolated, and other elements of [[Cellular respiration|cell respiration]] were determined to occur in the mitochondria.<ref name="Ernster-1981"/>

The first high-resolution electron [[micrographs]] appeared in 1952, replacing the Janus Green stains as the preferred way to visualize mitochondria.<ref name="Ernster-1981"/> This led to a more detailed analysis of the structure of the mitochondria, including confirmation that they were surrounded by a membrane. It also showed a second membrane inside the mitochondria that folded up in ridges dividing up the inner chamber and that the size and shape of the mitochondria varied from cell to cell.

The popular term "powerhouse of the cell" was coined by [[Philip Siekevitz]] in 1957.<ref name="Siekevitz-1957">{{cite journal |vauthors=Siekevitz P |title=Powerhouse of the cell |journal=[[Scientific American]] |year=1957 |volume=197 |pages=131–140|doi=10.1038/scientificamerican0757-131 |issue=1|bibcode=1957SciAm.197a.131S }}</ref><ref>{{cite journal | vauthors = Milane L, Trivedi M, Singh A, Talekar M, Amiji M | title = Mitochondrial biology, targets, and drug delivery | journal = Journal of Controlled Release | volume = 207 | pages = 40–58 | date = June 2015 | pmid = 25841699 | doi = 10.1016/j.jconrel.2015.03.036 }}</ref>

In 1967, it was discovered that mitochondria contained [[ribosomes]].<ref>{{cite journal | vauthors = Martin WF, Garg S, Zimorski V | title = Endosymbiotic theories for eukaryote origin | journal = Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume = 370 | issue = 1678 | pages = 20140330 | date = September 2015 | pmid = 26323761 | pmc = 4571569 | doi = 10.1098/rstb.2014.0330 }}</ref> In 1968, methods were developed for mapping the mitochondrial genes, with the genetic and physical map of yeast mitochondrial DNA completed in 1976.<ref name="Ernster-1981"/> In November 2024, Researchers from the United States have discovered that mitochondria divide into two distinct forms when cells are starved, this could help explain and describe how cancers thrive in hostile conditions.<ref>{{cite journal |last1=Thompson |first1=Benjamin |last2=Bates |first2=Emily |title=Surprise finding reveals mitochondrial 'energy factories' come in two different types |journal=Nature |date=6 November 2024 |doi=10.1038/d41586-024-03646-1 |pmid=39506172 }}</ref>