Editing Vesicle (biology and chemistry) (section)

==Types of vesicular structures==
[[Image:Hemozoin in food vacuole.jpg|thumb|250px|Electron micrograph of a cell containing a food vacuole (fv) and transport vacuole (tv) in a [[malaria|malaria parasite]] ]]

===Vacuoles===
[[Vacuoles]] are cellular organelles that contain mostly water.{{citation needed|date=March 2023}}
* [[Plant cells]] have a ''large central vacuole'' in the center of the cell that is used for [[osmoregulation|osmotic control]] and [[plant nutrition|nutrient]] storage.
* [[Contractile vacuole]]s are found in certain [[protists]], especially those in Phylum ''[[Ciliophora]]''. These vacuoles take water from the cytoplasm and excrete it from the cell to avoid bursting due to [[osmotic pressure]].

===Lysosomes===
* [[Lysosome]]s are involved in cellular digestion. Food can be taken from outside the cell into food vacuoles by a process called [[endocytosis]]. These food vacuoles fuse with lysosomes which break down the components so that they can be used in the cell. This form of cellular eating is called [[phagocytosis]].
* Lysosomes are also used to destroy defective or damaged organelles in a process called [[autophagy]]. They fuse with the membrane of the damaged organelle, digesting it.

===Transport vesicles===
* Transport vesicles can move molecules between locations inside the cell, e.g., proteins from the rough [[endoplasmic reticulum]] to the [[Golgi apparatus]].
* Membrane-bound and secreted proteins are made on [[ribosomes]] found in the [[rough endoplasmic reticulum]]. Most of these proteins mature in the [[Golgi apparatus]] before going to their final destination which may be to [[lysosomes]], [[peroxisomes]], or outside of the cell. These proteins travel within the cell inside transport vesicles.

===Secretory vesicles===
[[Secretory vesicles]] contain materials that are to be excreted from the cell. Cells have many reasons to excrete materials.
One reason is to dispose of wastes.
Another reason is tied to the function of the cell. Within a larger organism, some cells are specialized to produce certain chemicals. These chemicals are stored in secretory vesicles and released when needed.

====Types====
* [[Synaptic vesicle]]s are located at [[presynaptic terminal]]s in [[neuron]]s and store [[neurotransmitter]]s call [[Quantal neurotransmitter release|quanta]]. When a signal comes down an [[axon]], the synaptic vesicles fuse with the cell membrane releasing the neurotransmitter so that it can be detected by [[receptor (biochemistry)|receptor]] molecules on the next nerve cell.
* In animals, [[endocrine system|endocrine tissues]] release [[hormones]] into the bloodstream. These hormones are stored within secretory vesicles. A good example is an endocrine tissue found in the [[islets of Langerhans]] in the [[pancreas]]. This [[tissue (biology)|tissue]] contains many cell types that are defined by which hormones they produce.
* Secretory vesicles hold the enzymes that are used to make the [[cell wall]]s of [[plant cell|plants]], [[protist]]s, [[fungi]], [[bacteria]] and [[archaea]] cells as well as the [[extracellular matrix]] of [[animal cell]]s.
* Bacteria, [[archaea]], fungi and parasites release membrane vesicles (MVs) containing varied but specialized toxic compounds and biochemical signal molecules, which are transported to target cells to initiate processes in favour of the microbe, which include invasion of host cells and killing of competing microbes in the same niche.<ref name="DeatherageCookson2012">{{cite journal | vauthors = Deatherage BL, Cookson BT | title = Membrane vesicle release in bacteria, eukaryotes, and archaea: a conserved yet underappreciated aspect of microbial life | journal = Infection and Immunity | volume = 80 | issue = 6 | pages = 1948–57 | date = June 2012 | pmid = 22409932 | pmc = 3370574 | doi = 10.1128/IAI.06014-11 }}</ref>

===Extracellular vesicles===
[[Extracellular vesicles]] (EVs) are lipid bilayer-delimited particles produced by all domains of life including complex eukaryotes, both Gram-negative and Gram-positive bacteria, mycobacteria, and fungi.<ref name="Bible">{{cite journal | vauthors = Yáñez-Mó M, Siljander PR, Andreu Z, Zavec AB, Borràs FE, Buzas EI, Buzas K, Casal E, Cappello F, Carvalho J, Colás E, Cordeiro-da Silva A, Fais S, Falcon-Perez JM, Ghobrial IM, Giebel B, Gimona M, Graner M, Gursel I, Gursel M, Heegaard NH, Hendrix A, Kierulf P, Kokubun K, Kosanovic M, Kralj-Iglic V, Krämer-Albers EM, Laitinen S, Lässer C, Lener T, Ligeti E, Linē A, Lipps G, Llorente A, Lötvall J, Manček-Keber M, Marcilla A, Mittelbrunn M, Nazarenko I, Nolte-'t Hoen EN, Nyman TA, O'Driscoll L, Olivan M, Oliveira C, Pállinger É, Del Portillo HA, Reventós J, Rigau M, Rohde E, Sammar M, Sánchez-Madrid F, Santarém N, Schallmoser K, Ostenfeld MS, Stoorvogel W, Stukelj R, Van der Grein SG, Vasconcelos MH, Wauben MH, De Wever O | display-authors = 6 | title = Biological properties of extracellular vesicles and their physiological functions | journal = Journal of Extracellular Vesicles | volume = 4 | pages = 27066 | date = 2015 | pmid = 25979354 | pmc = 4433489 | doi = 10.3402/jev.v4.27066 }}</ref><ref name="MISEV2018">{{cite journal | vauthors = Théry C, Witwer KW, Aikawa E, Alcaraz MJ, Anderson JD, Andriantsitohaina R, Antoniou A, Arab T, Archer F, Atkin-Smith GK, Ayre DC, Bach JM, Bachurski D, Baharvand H, Balaj L, Baldacchino S, Bauer NN, Baxter AA, Bebawy M, Beckham C, Bedina Zavec A, Benmoussa A, Berardi AC, Bergese P, Bielska E, Blenkiron C, Bobis-Wozowicz S, Boilard E, Boireau W, Bongiovanni A, Borràs FE, Bosch S, Boulanger CM, Breakefield X, Breglio AM, Brennan MÁ, Brigstock DR, Brisson A, Broekman ML, Bromberg JF, Bryl-Górecka P, Buch S, Buck AH, Burger D, Busatto S, Buschmann D, Bussolati B, Buzás EI, Byrd JB, Camussi G, Carter DR, Caruso S, Chamley LW, Chang YT, Chen C, Chen S, Cheng L, Chin AR, Clayton A, Clerici SP, Cocks A, Cocucci E, Coffey RJ, Cordeiro-da-Silva A, Couch Y, Coumans FA, Coyle B, Crescitelli R, Criado MF, D'Souza-Schorey C, Das S, Datta Chaudhuri A, de Candia P, De Santana EF, De Wever O, Del Portillo HA, Demaret T, Deville S, Devitt A, Dhondt B, Di Vizio D, Dieterich LC, Dolo V, Dominguez Rubio AP, Dominici M, Dourado MR, Driedonks TA, Duarte FV, Duncan HM, Eichenberger RM, Ekström K, El Andaloussi S, Elie-Caille C, Erdbrügger U, Falcón-Pérez JM, Fatima F, Fish JE, Flores-Bellver M, Försönits A, Frelet-Barrand A, Fricke F, Fuhrmann G, Gabrielsson S, Gámez-Valero A, Gardiner C, Gärtner K, Gaudin R, Gho YS, Giebel B, Gilbert C, Gimona M, Giusti I, Goberdhan DC, Görgens A, Gorski SM, Greening DW, Gross JC, Gualerzi A, Gupta GN, Gustafson D, Handberg A, Haraszti RA, Harrison P, Hegyesi H, Hendrix A, Hill AF, Hochberg FH, Hoffmann KF, Holder B, Holthofer H, Hosseinkhani B, Hu G, Huang Y, Huber V, Hunt S, Ibrahim AG, Ikezu T, Inal JM, Isin M, Ivanova A, Jackson HK, Jacobsen S, Jay SM, Jayachandran M, Jenster G, Jiang L, Johnson SM, Jones JC, Jong A, Jovanovic-Talisman T, Jung S, Kalluri R, Kano SI, Kaur S, Kawamura Y, Keller ET, Khamari D, Khomyakova E, Khvorova A, Kierulf P, Kim KP, Kislinger T, Klingeborn M, Klinke DJ, Kornek M, Kosanović MM, Kovács ÁF, Krämer-Albers EM, Krasemann S, Krause M, Kurochkin IV, Kusuma GD, Kuypers S, Laitinen S, Langevin SM, Languino LR, Lannigan J, Lässer C, Laurent LC, Lavieu G, Lázaro-Ibáñez E, Le Lay S, Lee MS, Lee YX, Lemos DS, Lenassi M, Leszczynska A, Li IT, Liao K, Libregts SF, Ligeti E, Lim R, Lim SK, Linē A, Linnemannstöns K, Llorente A, Lombard CA, Lorenowicz MJ, Lörincz ÁM, Lötvall J, Lovett J, Lowry MC, Loyer X, Lu Q, Lukomska B, Lunavat TR, Maas SL, Malhi H, Marcilla A, Mariani J, Mariscal J, Martens-Uzunova ES, Martin-Jaular L, Martinez MC, Martins VR, Mathieu M, Mathivanan S, Maugeri M, McGinnis LK, McVey MJ, Meckes DG, Meehan KL, Mertens I, Minciacchi VR, Möller A, Møller Jørgensen M, Morales-Kastresana A, Morhayim J, Mullier F, Muraca M, Musante L, Mussack V, Muth DC, Myburgh KH, Najrana T, Nawaz M, Nazarenko I, Nejsum P, Neri C, Neri T, Nieuwland R, Nimrichter L, Nolan JP, Nolte-'t Hoen EN, Noren Hooten N, O'Driscoll L, O'Grady T, O'Loghlen A, Ochiya T, Olivier M, Ortiz A, Ortiz LA, Osteikoetxea X, Østergaard O, Ostrowski M, Park J, Pegtel DM, Peinado H, Perut F, Pfaffl MW, Phinney DG, Pieters BC, Pink RC, Pisetsky DS, Pogge von Strandmann E, Polakovicova I, Poon IK, Powell BH, Prada I, Pulliam L, Quesenberry P, Radeghieri A, Raffai RL, Raimondo S, Rak J, Ramirez MI, Raposo G, Rayyan MS, Regev-Rudzki N, Ricklefs FL, Robbins PD, Roberts DD, Rodrigues SC, Rohde E, Rome S, Rouschop KM, Rughetti A, Russell AE, Saá P, Sahoo S, Salas-Huenuleo E, Sánchez C, Saugstad JA, Saul MJ, Schiffelers RM, Schneider R, Schøyen TH, Scott A, Shahaj E, Sharma S, Shatnyeva O, Shekari F, Shelke GV, Shetty AK, Shiba K, Siljander PR, Silva AM, Skowronek A, Snyder OL, Soares RP, Sódar BW, Soekmadji C, Sotillo J, Stahl PD, Stoorvogel W, Stott SL, Strasser EF, Swift S, Tahara H, Tewari M, Timms K, Tiwari S, Tixeira R, Tkach M, Toh WS, Tomasini R, Torrecilhas AC, Tosar JP, Toxavidis V, Urbanelli L, Vader P, van Balkom BW, van der Grein SG, Van Deun J, van Herwijnen MJ, Van Keuren-Jensen K, van Niel G, van Royen ME, van Wijnen AJ, Vasconcelos MH, Vechetti IJ, Veit TD, Vella LJ, Velot É, Verweij FJ, Vestad B, Viñas JL, Visnovitz T, Vukman KV, Wahlgren J, Watson DC, Wauben MH, Weaver A, Webber JP, Weber V, Wehman AM, Weiss DJ, Welsh JA, Wendt S, Wheelock AM, Wiener Z, Witte L, Wolfram J, Xagorari A, Xander P, Xu J, Yan X, Yáñez-Mó M, Yin H, Yuana Y, Zappulli V, Zarubova J, Žėkas V, Zhang JY, Zhao Z, Zheng L, Zheutlin AR, Zickler AM, Zimmermann P, Zivkovic AM, Zocco D, Zuba-Surma EK | display-authors = 6 | title = Minimal information for studies of extracellular vesicles 2018 (MISEV2018): a position statement of the International Society for Extracellular Vesicles and update of the MISEV2014 guidelines | journal = Journal of Extracellular Vesicles | volume = 7 | issue = 1 | pages = 1535750 | date = 2018 | pmid = 30637094 | pmc = 6322352 | doi = 10.1080/20013078.2018.1535750 }}</ref>

==== Types ====
* [[Microvesicles|Ectosomes/microvesicles]] are shed directly from the plasma membrane and can range in size from around 30 [[Nanometre|nm]] to larger than a micron in diameter<ref name="vdPol2012" />{{rp|Table 1}}). These may include large particles such as [[bleb (cell biology)#Apoptotic function|apoptotic blebs]] released by dying cells,<ref name="J of T">{{cite journal | vauthors = van der Pol E, Böing AN, Gool EL, Nieuwland R | title = Recent developments in the nomenclature, presence, isolation, detection and clinical impact of extracellular vesicles | journal = Journal of Thrombosis and Haemostasis | volume = 14 | issue = 1 | pages = 48–56 | date = January 2016 | pmid = 26564379 | doi = 10.1111/jth.13190 | doi-access = free }}</ref><ref name="vdPol2012" />{{rp|Table 1}} [[large oncosomes]] released by some cancer cells, or "[[exophers]]," released by nematode neurons<ref name="Melentijevic-2017">{{cite journal | vauthors = Melentijevic I, Toth ML, Arnold ML, Guasp RJ, Harinath G, Nguyen KC, Taub D, Parker JA, Neri C, Gabel CV, Hall DH, Driscoll M | display-authors = 6 | title = C. elegans neurons jettison protein aggregates and mitochondria under neurotoxic stress | journal = Nature | volume = 542 | issue = 7641 | pages = 367–371 | date = February 2017 | pmid = 28178240 | pmc = 5336134 | doi = 10.1038/nature21362 | bibcode = 2017Natur.542..367M }}</ref> and mouse cardiomyocytes.
* [[Exosome (vesicle)|Exosome]]s: membranous vesicles of endocytic origin (30-100&nbsp;nm diameter).<ref name="vdPol2012" />{{rp|Table 1}}

Different types of EVs may be separated based on density<ref name=vdPol2012/>{{rp|Table 1}} (by gradient [[differential centrifugation]]), size, or surface markers.<ref>{{cite journal | vauthors = Mateescu B, Kowal EJ, van Balkom BW, Bartel S, Bhattacharyya SN, Buzás EI, Buck AH, de Candia P, Chow FW, Das S, Driedonks TA, Fernández-Messina L, Haderk F, Hill AF, Jones JC, Van Keuren-Jensen KR, Lai CP, Lässer C, Liegro ID, Lunavat TR, Lorenowicz MJ, Maas SL, Mäger I, Mittelbrunn M, Momma S, Mukherjee K, Nawaz M, Pegtel DM, Pfaffl MW, Schiffelers RM, Tahara H, Théry C, Tosar JP, Wauben MH, Witwer KW, Nolte-'t Hoen EN | display-authors = 6 | title = Obstacles and opportunities in the functional analysis of extracellular vesicle RNA - an ISEV position paper | journal = Journal of Extracellular Vesicles | volume = 6 | issue = 1 | pages = 1286095 | date = 2017 | pmid = 28326170 | pmc = 5345583 | doi = 10.1080/20013078.2017.1286095 }}</ref> However, EV subtypes have an overlapping size and density ranges, and subtype-unique markers must be established on a cell-by-cell basis. Therefore, it is difficult to pinpoint the biogenesis pathway that gave rise to a particular EV after it has left the cell.<ref name="MISEV2018" />

In humans, endogenous extracellular vesicles likely play a role in coagulation, intercellular signaling and waste management.<ref name=vdPol2012>{{cite journal | vauthors = van der Pol E, Böing AN, Harrison P, Sturk A, Nieuwland R | title = Classification, functions, and clinical relevance of extracellular vesicles | journal = Pharmacological Reviews | volume = 64 | issue = 3 | pages = 676–705 | date = July 2012 | pmid = 22722893 | doi = 10.1124/pr.112.005983 | s2cid = 7764903 }} Free full text</ref> They are also implicated in the pathophysiological processes involved in multiple diseases, including cancer.<ref>{{cite journal | vauthors = Dhondt B, Rousseau Q, De Wever O, Hendrix A | title = Function of extracellular vesicle-associated miRNAs in metastasis | journal = Cell and Tissue Research | volume = 365 | issue = 3 | pages = 621–41 | date = September 2016 | pmid = 27289232 | doi = 10.1007/s00441-016-2430-x | s2cid = 2746182 | hdl = 1854/LU-7250365 | url = https://figshare.com/articles/journal_contribution/7067858 }}</ref> Extracellular vesicles have raised interest as a potential source of biomarker discovery because of their role in intercellular communication, release into easily accessible body fluids and the resemblance of their molecular content to that of the releasing cells.<ref>{{cite journal | vauthors = Dhondt B, Van Deun J, Vermaerke S, de Marco A, Lumen N, De Wever O, Hendrix A | title = Urinary extracellular vesicle biomarkers in urological cancers: From discovery towards clinical implementation | journal = The International Journal of Biochemistry & Cell Biology | volume = 99 | pages = 236–256 | date = June 2018 | pmid = 29654900 | doi = 10.1016/j.biocel.2018.04.009 | hdl = 1854/LU-8559155 | s2cid = 4876604 | url = https://figshare.com/articles/journal_contribution/7067903 }}</ref> The extracellular vesicles of [[Mesenchymal stem cell|(mesenchymal) stem cells]], also known as the [[Stem cell secretome|secretome of stem cells]], are being researched and applied for therapeutic purposes, predominantly [[Degenerative disease|degenerative]], [[Autoimmune disease|auto-immune]] and/or [[Immune-mediated inflammatory diseases|inflammatory]] diseases.<ref>{{cite journal | vauthors = Teixeira FG, Carvalho MM, Sousa N, Salgado AJ | title = Mesenchymal stem cells secretome: a new paradigm for central nervous system regeneration? | journal = Cellular and Molecular Life Sciences | volume = 70 | issue = 20 | pages = 3871–82 | date = October 2013 | pmid = 23456256 | doi = 10.1007/s00018-013-1290-8 | pmc = 11113366 | s2cid = 18640402 | hdl = 1822/25128 | hdl-access = free }}</ref>

In Gram-negative bacteria, EVs are produced by the pinching off of the outer membrane; however, how EVs escape the thick cell walls of Gram-positive bacteria, mycobacteria and fungi is still unknown. These EVs contain varied cargo, including nucleic acids, toxins, lipoproteins and enzymes and have important roles in microbial physiology and pathogenesis. In host–pathogen interactions, gram negative bacteria produce vesicles which play roles in establishing a colonization niche, carrying and transmitting virulence factors into host cells and modulating host defense and response.<ref name=Kuehn2005>{{cite journal | vauthors = Kuehn MJ, Kesty NC | title = Bacterial outer membrane vesicles and the host–pathogen interaction | journal = Genes & Development | volume = 19 | issue = 22 | pages = 2645–55 | date = November 2005 | pmid = 16291643 | doi = 10.1101/gad.1299905 | doi-access = free }}</ref>

Ocean [[cyanobacteria]] have been found to continuously release vesicles containing proteins, DNA and RNA into the open ocean. Vesicles carrying DNA from diverse bacteria are abundant in coastal and open-ocean seawater samples.<ref name=Biller2014>{{cite journal | vauthors = Biller SJ, Schubotz F, Roggensack SE, Thompson AW, Summons RE, Chisholm SW | title = Bacterial vesicles in marine ecosystems | journal = Science | volume = 343 | issue = 6167 | pages = 183–6 | date = January 2014 | pmid = 24408433 | doi = 10.1126/science.1243457 | s2cid = 206551356 | bibcode = 2014Sci...343..183B | hdl = 1721.1/84545 | hdl-access = free }}</ref>

===Protocells===
The [[RNA world]] hypothesis assumes that the first [[self-replication|self-replicating]] [[genome]]s were strands of RNA.  This hypothesis contains the idea that RNA strands formed [[ribozymes]] (folded RNA molecules) capable of catalyzing RNA replication.  These primordial biological catalysis were considered to be contained within vesicles ([[protocell]]s) with membranes composed of [[fatty acid]]s and related [[amphiphile]]s.<ref name="Adamala2013">{{cite journal |vauthors=Adamala K, Szostak JW |title=Nonenzymatic template-directed RNA synthesis inside model protocells |journal=Science |volume=342 |issue=6162 |pages=1098–1100 |date=November 2013 |pmid=24288333 |pmc=4104020 |doi=10.1126/science.1241888 |bibcode=2013Sci...342.1098A }}</ref>  Template-directed RNA synthesis by the copying of RNA templates inside fatty acid vesicles has been demonstrated by Adamata and Szostak.<ref name = Adamala2013/>

===Other types===
{{Main|Gas vesicle}}

Gas vesicles are used by [[archaea]], [[bacteria]] and [[plankton]]ic microorganisms, possibly to control vertical migration by regulating the gas content and thereby [[buoyancy]], or possibly to position the cell for maximum solar light harvesting.  These vesicles are typically lemon-shaped or cylindrical tubes made out of protein;<ref name=Pfeifer2012>{{cite journal | vauthors = Pfeifer F | title = Distribution, formation and regulation of gas vesicles | journal = Nature Reviews. Microbiology | volume = 10 | issue = 10 | pages = 705–15 | date = October 2012 | pmid = 22941504 | doi = 10.1038/nrmicro2834 | s2cid = 9926129 }}</ref> their diameter determines the strength of the vesicle with larger ones being weaker. The diameter of the vesicle also affects its volume and how efficiently it can provide buoyancy.  In cyanobacteria, natural selection has worked to create vesicles that are at the maximum diameter possible while still being structurally stable.  The protein skin is permeable to gases but not water, keeping the vesicles from flooding.<ref name="Gas vesicles"/>

[[Extracellular matrix|Matrix]] vesicles are located within the extracellular space, or matrix. Using [[electron microscopy]], they were discovered independently in 1967 by H. Clarke Anderson<ref>{{cite journal | vauthors = Anderson HC | title = Electron microscopic studies of induced cartilage development and calcification | journal = The Journal of Cell Biology | volume = 35 | issue = 1 | pages = 81–101 | date = October 1967 | pmid = 6061727 | pmc = 2107116 | doi = 10.1083/jcb.35.1.81 }}</ref> and Ermanno Bonucci.<ref>{{cite journal | vauthors = Bonucci E | title = Fine structure of early cartilage calcification | journal = Journal of Ultrastructure Research | volume = 20 | issue = 1 | pages = 33–50 | date = September 1967 | pmid = 4195919 | doi = 10.1016/S0022-5320(67)80034-0 }}</ref> These cell-derived vesicles are specialized to initiate [[biomineralisation]] of the matrix in a variety of tissues, including [[bone]], [[cartilage]] and [[dentin]]. During normal [[calcification]], a major influx of calcium and phosphate ions into the cells accompanies cellular [[apoptosis]] (genetically determined self-destruction) and matrix vesicle formation. Calcium-loading also leads to formation of [[phosphatidylserine]]:calcium:phosphate complexes in the plasma membrane mediated in part by a protein called [[annexins]]. Matrix vesicles bud from the plasma membrane at sites of interaction with the extracellular matrix. Thus, matrix vesicles convey to the extracellular matrix calcium, phosphate, lipids and the annexins which act to nucleate mineral formation. These processes are precisely coordinated to bring about, at the proper place and time, mineralization of the tissue's matrix unless the Golgi are non-existent.{{citation needed|date=March 2023}}

[[Endosome|Multivesicular body]], or MVB, is a membrane-bound vesicle containing a number of smaller vesicles.<ref>{{Cite journal |last1=Von Bartheld |first1=Christopher S. |last2=Altick |first2=Amy L. |date=March 2011 |title=Multivesicular bodies in neurons: Distribution, protein content, and trafficking functions |journal=Progress in Neurobiology |language=en |volume=93 |issue=3 |pages=313–340 |doi=10.1016/j.pneurobio.2011.01.003|pmid=21216273 |pmc=3055956 }}</ref>