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Endoplasmic reticulum

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777:. The only structural difference between this organelle and the smooth endoplasmic reticulum is the composition of proteins they have, both bound to their membranes and drifting within the confines of their lumens. This fundamental difference is indicative of their functions: The endoplasmic reticulum synthesizes molecules, while the sarcoplasmic reticulum stores calcium ions and pumps them out into the sarcoplasm when the muscle fiber is stimulated. After their release from the sarcoplasmic reticulum, calcium ions interact with contractile proteins that utilize ATP to shorten the muscle fiber. The sarcoplasmic reticulum plays a major role in 672: 416: 443:. The functions of the endoplasmic reticulum can be summarized as the synthesis and export of proteins and membrane lipids, but varies between ER and cell type and cell function. The quantity of both rough and smooth endoplasmic reticulum in a cell can slowly interchange from one type to the other, depending on the changing metabolic activities of the cell. Transformation can include embedding of new proteins in membrane as well as structural changes. Changes in protein content may occur without noticeable structural changes. 747: 148: 916: 315: 53: 683:) is scarce. Instead there are areas where the ER is partly smooth and partly rough, this area is called the transitional ER. The transitional ER gets its name because it contains ER exit sites. These are areas where the transport vesicles which contain lipids and proteins made in the ER, detach from the ER and start moving to the 483:. However, the ribosomes are not a stable part of this organelle's structure as they are constantly being bound and released from the membrane. A ribosome only binds to the RER once a specific protein-nucleic acid complex forms in the cytosol. This special complex forms when a free ribosome begins 737:
and consists of tubules that are located near the cell periphery. These tubes sometimes branch forming a network that is reticular in appearance. In some cells, there are dilated areas like the sacs of rough endoplasmic reticulum. The network of smooth endoplasmic reticulum allows for an increased
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Increased and supraphysiological ER stress in pancreatic β cells disrupts normal insulin secretion, leading to hyperinsulinemia and consequently peripheral insulin resistance associated with obesity in humans. Human clinical trials also suggested a causal link between obesity-induced increase in
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Klein, Marie-Christine; Zimmermann, Katharina; Schorr, Stefan; Landini, Martina; Klemens, Patrick A. W.; Altensell, Jacqueline; Jung, Martin; Krause, Elmar; Nguyen, Duy; Helms, Volkhard; Rettig, Jens; Fecher-Trost, Claudia; Cavalié, Adolfo; Hoth, Markus; Bogeski, Ivan; Neuhaus, H. Ekkehard;
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Moreno JA, Halliday M, Molloy C, Radford H, Verity N, Axten JM, Ortori CA, Willis AE, Fischer PM, Barrett DA, Mallucci GR (October 2013). "Oral treatment targeting the unfolded protein response prevents neurodegeneration and clinical disease in prion-infected mice".
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The endoplasmic reticulum does not harbor an ATP-regeneration machinery, and therefore requires ATP import from mitochondria. The imported ATP is vital for the ER to carry out its house keeping cellular functions, such as for protein folding and trafficking.
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Kannan, Muthukumar; Riekhof, Wayne R.; Voelker, Dennis R. (February 2015). "Transport of Phosphatidylserine from the Endoplasmic Reticulum to the Site of Phosphatidylserine Decarboxylase2 in Yeast: Phosphatidylserine Transport to the Locus of Psd2p".
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for transmembrane proteins. However, variations of KDEL and KKXX do occur, and other sequences can also give rise to endoplasmic reticulum retention. It is not known whether such variation can lead to sub-ER localizations. There are three KDEL
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have an abundance of smooth endoplasmic reticulum. It also carries out the metabolism of carbohydrates, detoxification of natural metabolism products and of alcohol and drugs, attachment of receptors on cell membrane proteins, and
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toward their destination. In human fibroblasts, the ER is always co-distributed with microtubules and the depolymerisation of the latter cause its co-aggregation with mitochondria, which are also associated with the ER.
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The endoplasmic reticulum is also part of a protein sorting pathway. It is, in essence, the transportation system of the eukaryotic cell. The majority of its resident proteins are retained within it through a retention
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Ozcan U, Cao Q, Yilmaz E, Lee AH, Iwakoshi NN, Ozdelen E, Tuncman G, Görgün C, Glimcher LH, Hotamisligil GS (October 2004). "Endoplasmic reticulum stress links obesity, insulin action, and type 2 diabetes".
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concentration. Smooth endoplasmic reticulum is found in a variety of cell types (both animal and plant), and it serves different functions in each. The smooth endoplasmic reticulum also contains the enzyme
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contain different ratios of the two types of ER depending on the activities of the cell. RER is found mainly toward the nucleus of cell and SER towards the cell membrane or plasma membrane of cell.
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Görlich D, Prehn S, Hartmann E, Kalies KU, Rapoport TA (October 1992). "A mammalian homolog of SEC61p and SECYp is associated with ribosomes and nascent polypeptides during translocation".
519:), which removes the signal peptide. Ribosomes at this point may be released back into the cytosol; however, non-translating ribosomes are also known to stay associated with translocons. 452: 2229:"KDEL and KKXX retrieval signals appended to the same reporter protein determine different trafficking between endoplasmic reticulum, intermediate compartment, and Golgi complex" 853:, are moved across the endoplasmic reticulum membrane. Proteins that are transported by the endoplasmic reticulum throughout the cell are marked with an address tag called a 2192:
Soltys BJ, Gupta RS (1992). "Interrelationships of endoplasmic reticulum, mitochondria, intermediate filaments, and microtubules--a quadruple fluorescence labeling study".
1042: 526:. The double membrane sheets are stacked and connected through several right- or left-handed helical ramps, the "Terasaki ramps", giving rise to a structure resembling a 2583:
van Vliet, Stephan; Koh, Han-Chow E.; Patterson, Bruce W.; Yoshino, Mihoko; LaForest, Richard; Gropler, Robert J.; Klein, Samuel; Mittendorfer, Bettina (1 October 2020).
907:) receptors in mammalian cells, and they have a very high degree of sequence identity. The functional differences between these receptors remain to be established. 522:
The membrane of the rough endoplasmic reticulum is in the form of large double-membrane sheets that are located near, and continuous with, the outer layer of the
825:, and the peptidylprolyl isomerase family. Only properly folded proteins are transported from the rough ER to the Golgi apparatus – unfolded proteins cause an 801:. Rough endoplasmic reticulum is also involved in protein synthesis. Correct folding of newly made proteins is made possible by several endoplasmic reticulum 562:, where the membranes of the endoplasmic reticulum and other organelles are held closely together, allowing the transfer of lipids and other small molecules. 2327:"Translocation of ATP into the lumen of rough endoplasmic reticulum-derived vesicles and its binding to luminal proteins including BiP (GRP 78) and GRP 94" 1836:
Friedman, Jonathan R.; Kannan, Muthukumar; Toulmay, Alexandre; Jan, Calvin H.; Weissman, Jonathan S.; Prinz, William A.; Nunnari, Jodi (January 2018).
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Terasaki M, Shemesh T, Kasthuri N, Klemm RW, Schalek R, Hayworth KJ, Hand AR, Yankova M, Huber G, Lichtman JW, Rapoport TA, Kozlov MM (July 2013).
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mechanism shows sensitivity to cytosolic Ca ranging from high nM to low μM range, with the Ca-sensing element yet to be identified and validated.
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Kober L, Zehe C, Bode J (October 2012). "Development of a novel ER stress based selection system for the isolation of highly productive clones".
199:). The endoplasmic reticulum is found in most eukaryotic cells and forms an interconnected network of flattened, membrane-enclosed sacs known as 2730:
Kaser A, Lee AH, Franke A, Glickman JN, Zeissig S, Tilg H, Nieuwenhuis EE, Higgins DE, Schreiber S, Glimcher LH, Blumberg RS (September 2008).
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Toyoshima C, Nakasako M, Nomura H, Ogawa H (June 2000). "Crystal structure of the calcium pump of sarcoplasmic reticulum at 2.6 A resolution".
1997: 869:, a membrane-embedded multiprotein complex. Proteins that are destined for places outside the endoplasmic reticulum are packed into transport 459:
is synthesized and secreted into the rough endoplasmic reticulum, which appears at the upper right approximately halfway through the animation
2895: 17: 687:. Specialized cells can have a lot of smooth endoplasmic reticulum and in these cells the smooth ER has many functions. It synthesizes 2424:
Yong, Jing; Bischof, Helmut; Burgstaller, Sandra; Siirin, Marina; Murphy, Anne; Malli, Roland; Kaufman, Randal J (9 September 2019).
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that work as an address tag, which are removed when the polypeptide reaches its destination. Nascent peptides reach the ER via the
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The ER ATP transporter, SLC35B1/AXER, was recently cloned and characterized, and the mitochondria supply ATP to the ER through a
2585:"Obesity Is Associated With Increased Basal and Postprandial β-Cell Insulin Secretion Even in the Absence of Insulin Resistance" 1591: 1612:
Levine T (September 2004). "Short-range intracellular trafficking of small molecules across endoplasmic reticulum junctions".
1682:Čaval, Tomislav; Zhu, Jing; Tian, Weihua; Remmelzwaal, Sanne; Yang, Zhang; Clausen, Henrik; Heck, Albert J. R. (2019-01-01). 1375: 1281: 1248: 1112: 550:
marks them to be brought back to the rough endoplasmic reticulum. The rough endoplasmic reticulum works in concert with the
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giving it a "rough" appearance (hence its name). The binding site of the ribosome on the rough endoplasmic reticulum is the
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Yong, Jing; Bischof, Helmut; Burgstaller, Sandra; Siirin, Marina; Murphy, Anne; Malli, Roland; Kaufman, Randal J (2019).
886:. This motif is composed of four amino acids at the end of the protein sequence. The most common retention sequences are 539: 833:
regulation, calcium regulation, glucose deprivation, and viral infection or the over-expression of proteins can lead to
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to their proper destinations. The second method of transport out of the endoplasmic reticulum involves areas called
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related to the endoplasmic reticulum. The UPR is activated in response to an accumulation of unfolded or misfolded
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The endoplasmic reticulum serves many general functions, including the folding of protein molecules in sacs called
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Mittendorfer, Bettina; Patterson, Bruce W.; Smith, Gordon I.; Yoshino, Mihoko; Klein, Samuel (1 February 2022).
3312: 2732:"XBP1 links ER stress to intestinal inflammation and confers genetic risk for human inflammatory bowel disease" 814: 512: 3059: 2681:
Casas-Tinto S, Zhang Y, Sanchez-Garcia J, Gomez-Velazquez M, Rincon-Limas DE, Fernandez-Funez P (June 2011).
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Guven J, Huber G, Valencia DM (October 2014). "Terasaki spiral ramps in the rough endoplasmic reticulum".
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of the endoplasmic reticulum. The UPR functions to restore normal function of the cell by halting protein
1838:"Lipid Homeostasis Is Maintained by Dual Targeting of the Mitochondrial PE Biosynthesis Enzyme to the ER" 1019: 806: 504: 81: 959:
and subsequently causes a higher susceptibility for inflammatory processes that may even contribute to
891: 887: 599: 203:(in the RER), and tubular structures in the SER. The membranes of the ER are continuous with the outer 738:
surface area to be devoted to the action or storage of key enzymes and the products of these enzymes.
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Levine T, Loewen C (August 2006). "Inter-organelle membrane contact sites: through a glass, darkly".
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Kannan, Muthukumar; Lahiri, Sujoy; Liu, Li-Ka; Choudhary, Vineet; Prinz, William A. (March 2017).
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proteins, either secreted constitutively with no tag or secreted in a regulatory manner involving
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Raykhel I, Alanen H, Salo K, Jurvansuu J, Nguyen VD, Latva-Ranta M, Ruddock L (December 2007).
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Stornaiuolo M, Lotti LV, Borgese N, Torrisi MR, Mottola G, Martire G, Bonatti S (March 2003).
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Yong, Jing; Johnson, James D.; Arvan, Peter; Han, Jaeseok; Kaufman, Randal J. (August 2021).
1743:"Phosphatidylserine synthesis at membrane contact sites promotes its transport out of the ER" 722: 559: 484: 432: 306:, which means "network", was applied by Porter in 1953 to describe this fabric of membranes. 1297: 3291: 2380: 2150: 1954: 1542: 155:(shown in the lower right-hand area of the picture). Dark small circles in the network are 8: 2917: 995: 802: 726: 574: 289: 2384: 2154: 1958: 1546: 675:
Electron micrograph showing smooth ER (arrow) in mouse tissue, at 110,510× magnification
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The general structure of the endoplasmic reticulum is a network of membranes called
415: 2804: 2751: 2743: 2702: 2694: 2653: 2645: 2604: 2596: 2555: 2547: 2506: 2488: 2447: 2437: 2426:"Mitochondria supply ATP to the ER through a mechanism antagonized by cytosolic Ca" 2396: 2388: 2338: 2297: 2289: 2248: 2240: 2201: 2158: 2106: 2069: 2061: 1982: 1962: 1917: 1909: 1857: 1849: 1802: 1764: 1754: 1713: 1695: 1656: 1621: 1554: 1550: 1505: 1497: 1448: 1404: 1324: 1206: 1198: 1157: 1149: 1100: 910: 883: 734: 523: 302: 281: 204: 1486:"Stacked endoplasmic reticulum sheets are connected by helicoidal membrane motifs" 1065: 2942: 2868: 2808: 1853: 1822: 999: 987: 964: 798: 750: 730: 708: 684: 603: 531: 388: 293: 262: 176: 86: 3101: 2536:"Therapeutic opportunities for pancreatic β-cell ER stress in diabetes mellitus" 2905: 2861: 2747: 2551: 2392: 1501: 1453: 1436: 1238: 967:, XBP1 anomalies have been linked to the inflammatory bowel diseases including 854: 602:
that stay embedded in the membrane as vesicles exit and bind to new membranes.
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shuttle proteins between these two compartments. Vesicles are surrounded by
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and the inhibition of the UPR could become a treatment for those diseases.
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Ca2+-antagonized transport into the endoplasmic reticulum (CaATiER) model
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as assembly continues. This is N-linked (O-linking occurs in the Golgi).
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and engage in certain common activities such as the synthesis of certain
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encloses the cisternal space (or lumen), which is continuous with the
3286: 3248: 3221: 3193: 3086: 3069: 3004: 2977: 2909: 2278:"A molecular specificity code for the three mammalian KDEL receptors" 2065: 1966: 1913: 641: 585: 516: 288:. The lacy membranes of the endoplasmic reticulum were first seen by 180: 172: 120: 116: 60: 2205: 1187:"Observations on a submicroscopic basophilic component of cytoplasm" 507:(SRP). Translation pauses and the ribosome complex binds to the RER 3231: 2987: 2972: 2967: 2840: 2367:
Zimmermann, Richard; Lang, Sven; Haferkamp, Ilka (28 August 2018).
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Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002).
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The surface of the rough endoplasmic reticulum (often abbreviated
3253: 3153: 3106: 2982: 2631: 1582:"Answers - the Most Trusted Place for Answering Life's Questions" 983: 704: 653: 649: 607: 440: 367: 238: 219: 112: 107: 2683:"The ER stress factor XBP1s prevents amyloid-beta neurotoxicity" 911:
Bioenergetics regulation of ER ATP supply by a CaATiER mechanism
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Clairmont, CA; De Maio, A; Hirschberg, CB (25 February 1992).
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The rough endoplasmic reticulum is key in multiple functions:
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Buvat, R. (1963). "Electron Microscopy of Plant Protoplasm".
1003: 1002:. Sustained overactivation of the UPR has been implicated in 830: 810: 611: 543: 274: 270: 254: 223: 151:
Micrograph of rough endoplasmic reticulum network around the
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Cell organelle that synthesizes, folds and processes proteins
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In most cells the smooth endoplasmic reticulum (abbreviated
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A 2-minute animation showing how a protein destined for the
2793: 1579:. Retrieved September 13, 2006, from Answers.com Web site: 952: 699:. Cells which secrete these products, such as those in the 624:
N-linked glycosylation: If the protein is properly folded,
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Animations of the various cell functions referenced here
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Endoplasmic Reticulum- Structure, Types & Functions
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Lipid and protein composition of Endoplasmic reticulum
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insulin secretion and peripheral insulin resistance.
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The sarcoplasmic reticulum (SR), from the Greek σάρξ
427:. These sac-like structures are held together by the 1791: 1430: 1428: 2533: 2047: 249:. The RER is especially prominent in cells such as 1577:McGraw-Hill Encyclopedia of Science and Technology 1532: 1477: 1359: 1269: 1135: 753:fiber, with sarcoplasmic reticulum colored in blue 2139: 1435:Shibata Y, Voeltz GK, Rapoport TA (August 2006). 1425: 175:, and has many other important functions such as 3304: 829:as a stress response in the ER. Disturbances in 2048:Xu C, Bailly-Maitre B, Reed JC (October 2005). 1895: 1390: 666: 2781:"Peter Walter's Short Talk: Unfolding the UPR" 1232: 1230: 446: 269:. The SER is especially abundant in mammalian 171:) is a part of a transportation system of the 2889: 2096: 1136:Porter KR, Claude A, Fullam EF (March 1945). 793:and the transport of synthesized proteins in 2723: 2674: 2090: 2002:(3rd ed.). Academic Press. p. 69. 1938: 1646: 1276:(2nd ed.). Washington (DC): ASM Press. 606:proteins are key in targeting the membrane; 546:targets vesicles to the Golgi apparatus and 207:. The endoplasmic reticulum is not found in 2191: 1243:(4th ed.). New York: Garland Science. 1227: 1178: 1088: 253:. The SER lacks ribosomes and functions in 218:The two types of ER share many of the same 2896: 2882: 1384: 1097:International Review of Cytology Volume 14 1039:"Endoplasmic Reticulum (Rough and Smooth)" 2755: 2706: 2657: 2608: 2559: 2510: 2492: 2451: 2441: 2400: 2342: 2301: 2252: 2073: 2023:Martini F, Nath J, Bartholomew E (2014). 1921: 1861: 1768: 1758: 1717: 1699: 1509: 1452: 1408: 1210: 1161: 741: 1391:Seiser RM, Nicchitta CV (October 2000). 1366:(5th ed.). W. H. Freeman. pp.  942: 914: 745: 670: 475:) is studded with protein-manufacturing 450: 414: 348: Smooth endoplasmic reticulum (SER) 313: 300:, and Ernest F. Fullam. Later, the word 284:by Garnier in 1897, who coined the term 146: 1995: 1896:Maxfield FR, Wüstner D (October 2002). 1063: 861:chain (i.e., a protein) contains a few 404: Trans face of the Golgi apparatus 341: Rough endoplasmic reticulum (RER) 14: 3305: 2025:Fundamentals of Anatomy and Physiology 1611: 1357: 1351: 1267: 1184: 1129: 411: Cisternae of the Golgi apparatus 2877: 2054:The Journal of Clinical Investigation 1902:The Journal of Clinical Investigation 1898:"Intracellular cholesterol transport" 1526: 1094: 1045:from the original on 24 November 2015 957:endoplasmic reticulum stress response 835:endoplasmic reticulum stress response 614:proteins are key in the fusion event. 511:where translation continues with the 419:3D rendering of endoplasmic reticulum 397: Cis face of the Golgi apparatus 57:Components of a typical animal cell: 1191:The Journal of Experimental Medicine 1142:The Journal of Experimental Medicine 929:Ca-antagonized transport into the ER 844: 592:and paired basic amino acids in the 2331:The Journal of Biological Chemistry 1688:Molecular & Cellular Proteomics 1397:The Journal of Biological Chemistry 1041:. British Society of Cell Biology. 1031: 24: 2778: 1006:diseases as well as several other 241:) face of the RER is studded with 25: 3324: 2834: 2638:Journal of Clinical Investigation 1437:"Rough sheets and smooth tubules" 1099:. Vol. 14. pp. 41–155. 773:("flesh"), is smooth ER found in 2099:Biotechnology and Bioengineering 1257:from the original on 2017-10-03. 857:. The N-terminus (one end) of a 716:. In muscle cells, it regulates 648:-acetylglucosamine, 9-branching 51: 2787: 2772: 2625: 2576: 2527: 2468: 2417: 2359: 2318: 2269: 2220: 2185: 2133: 2041: 2016: 1989: 1889: 1878: 1829: 1785: 1734: 1675: 1649:Current Opinion in Cell Biology 1640: 1605: 1594:from the original on 2006-11-16 1575:Endoplasmic reticulum. (n.d.). 1569: 890:for lumen-located proteins and 779:excitation-contraction coupling 765:Calcium-induced calcium release 2797:Science Translational Medicine 1555:10.1103/PhysRevLett.113.188101 1358:Lodish H, et al. (2003). 1308: 1290: 1272:The cell: a molecular approach 1261: 1057: 656:at the end) to the side-chain 491:of a protein destined for the 473:granular endoplasmic reticulum 13: 1: 3060:Microtubule organizing center 2344:10.1016/S0021-9258(19)50622-6 2233:Molecular Biology of the Cell 2194:Biochemistry and Cell Biology 1240:Molecular biology of the cell 1105:10.1016/S0074-7696(08)60021-2 1025: 2809:10.1126/scitranslmed.3006767 2540:Nature Reviews Endocrinology 1854:10.1016/j.devcel.2017.11.023 1329:10.1016/0092-8674(92)90517-G 784: 667:Smooth endoplasmic reticulum 309: 193:smooth endoplasmic reticulum 98:Smooth endoplasmic reticulum 18:Smooth endoplasmic reticulum 7: 2282:The Journal of Cell Biology 1020:Ribosome-associated vesicle 1013: 849:Secretory proteins, mostly 807:protein disulfide isomerase 628:recognizes the AA sequence 505:signal recognition particle 447:Rough endoplasmic reticulum 185:rough endoplasmic reticulum 82:Rough endoplasmic reticulum 10: 3329: 2748:10.1016/j.cell.2008.07.021 2552:10.1038/s41574-021-00510-4 2393:10.1038/s41467-018-06003-9 2027:(10th ed.). Pearson. 1502:10.1016/j.cell.2013.06.031 1454:10.1016/j.cell.2006.07.019 1008:neurodegenerative diseases 762: 756: 600:Integral membrane proteins 183:made up of two subunits – 3279: 3214: 3139: 3030: 2916: 1996:Goodman SR (2007-11-26). 1747:Journal of Lipid Research 1661:10.1016/j.ceb.2006.06.011 1626:10.1016/j.tcb.2004.07.017 976:unfolded protein response 827:unfolded protein response 733:. It is connected to the 626:oligosaccharyltransferase 50: 41: 32: 3055:Prokaryotic cytoskeleton 2687:Human Molecular Genetics 1701:10.1074/mcp.RA118.000967 1070:Bibliographie Anatomique 980:cellular stress response 119:; with which, comprises 2245:10.1091/mbc.E02-08-0468 2163:10.1126/science.1103160 1535:Physical Review Letters 542:called COPI and COPII. 280:The ER was observed by 1614:Trends in Cell Biology 1410:10.1074/jbc.M004462200 1362:Molecular Cell Biology 1185:PORTER KR (May 1953). 920: 759:Sarcoplasmic reticulum 754: 742:Sarcoplasmic reticulum 729:to glucose, a step in 676: 560:membrane contact sites 460: 439:but separate from the 420: 412: 245:that are the sites of 160: 3313:Endoplasmic reticulum 3045:Intermediate filament 2938:Endoplasmic reticulum 2846:Endoplasmic Reticulum 2373:Nature Communications 2294:10.1083/jcb.200705180 955:lead to a heightened 943:Clinical significance 918: 749: 723:glucose-6-phosphatase 674: 499:polymerized encode a 454: 433:phospholipid membrane 418: 317: 230:. Different types of 165:endoplasmic reticulum 150: 115:(fluid that contains 3292:Extracellular matrix 1999:Medical Cell Biology 1203:10.1084/jem.97.5.727 1154:10.1084/jem.81.3.233 1064:Garnier, C. (1897). 873:and moved along the 805:proteins, including 577:marker added in the 370:that are transported 261:, the production of 2995:Cytoplasmic granule 2494:10.7554/eLife.49682 2443:10.7554/eLife.49682 2385:2018NatCo...9.3489K 2155:2004Sci...306..457O 1959:2000Natur.405..647T 1760:10.1194/jlr.M072959 1547:2014PhRvL.113r8101G 1302:The Free Dictionary 961:Alzheimer's disease 727:glucose-6-phosphate 575:mannose-6-phosphate 556:target new proteins 290:electron microscopy 73:(dots as part of 5) 44:Animal cell diagram 3020:Weibel–Palade body 2904:Structures of the 2867:2008-04-22 at the 2699:10.1093/hmg/ddr100 2200:(10–11): 1174–86. 1842:Developmental Cell 1268:Cooper GM (2000). 921: 809:(PDI), ERp29, the 755: 714:steroid metabolism 677: 536:transport vesicles 461: 421: 413: 257:synthesis but not 179:. It is a type of 161: 3300: 3299: 3080:Spindle pole body 2803:(206): 206ra138. 2650:10.1172/JCI154068 2601:10.2337/db20-0377 2595:(10): 2112–2119. 2111:10.1002/bit.24527 1848:(2): 261–270.e6. 1807:10.1111/tra.12236 1377:978-0-7167-4366-8 1283:978-0-87893-106-4 1250:978-0-8153-3218-3 1114:978-0-12-364314-8 951:Abnormalities in 935:) mechanism. The 845:Protein transport 839:unfolded proteins 725:, which converts 534:, membrane-bound 495:. The first 5–30 493:secretory pathway 457:secretory pathway 437:perinuclear space 247:protein synthesis 145: 144: 16:(Redirected from 3320: 2898: 2891: 2884: 2875: 2874: 2829: 2828: 2791: 2785: 2784: 2776: 2770: 2769: 2759: 2727: 2721: 2720: 2710: 2678: 2672: 2671: 2661: 2629: 2623: 2622: 2612: 2580: 2574: 2573: 2563: 2531: 2525: 2524: 2514: 2496: 2472: 2466: 2465: 2455: 2445: 2421: 2415: 2414: 2404: 2363: 2357: 2356: 2346: 2322: 2316: 2315: 2305: 2273: 2267: 2266: 2256: 2224: 2218: 2217: 2189: 2183: 2182: 2149:(5695): 457–61. 2137: 2131: 2130: 2105:(10): 2599–611. 2094: 2088: 2087: 2077: 2066:10.1172/JCI26373 2045: 2039: 2038: 2020: 2014: 2013: 1993: 1987: 1986: 1967:10.1038/35015017 1953:(6787): 647–55. 1942: 1936: 1935: 1925: 1914:10.1172/JCI16500 1893: 1887: 1886: 1882: 1876: 1875: 1865: 1833: 1827: 1826: 1789: 1783: 1782: 1772: 1762: 1738: 1732: 1731: 1721: 1703: 1679: 1673: 1672: 1644: 1638: 1637: 1609: 1603: 1602: 1600: 1599: 1573: 1567: 1566: 1530: 1524: 1523: 1513: 1481: 1475: 1474: 1456: 1432: 1423: 1422: 1412: 1388: 1382: 1381: 1365: 1355: 1349: 1348: 1312: 1306: 1305: 1294: 1288: 1287: 1275: 1265: 1259: 1258: 1234: 1225: 1224: 1214: 1182: 1176: 1175: 1165: 1133: 1127: 1126: 1092: 1086: 1085: 1061: 1055: 1054: 1052: 1050: 1035: 735:nuclear envelope 709:sebaceous glands 540:coating proteins 524:nuclear envelope 407: 400: 393: 383: 377: Transport 373: 362: 351: 344: 337: 327: 282:light microscopy 263:steroid hormones 205:nuclear membrane 89:(or, Golgi body) 55: 45: 36: 30: 29: 21: 3328: 3327: 3323: 3322: 3321: 3319: 3318: 3317: 3303: 3302: 3301: 3296: 3275: 3210: 3135: 3026: 2943:Golgi apparatus 2919: 2912: 2902: 2869:Wayback Machine 2837: 2832: 2792: 2788: 2779:Walter, Peter. 2777: 2773: 2728: 2724: 2693:(11): 2144–60. 2679: 2675: 2630: 2626: 2581: 2577: 2532: 2528: 2473: 2469: 2422: 2418: 2364: 2360: 2323: 2319: 2288:(6): 1193–204. 2274: 2270: 2225: 2221: 2206:10.1139/o92-163 2190: 2186: 2138: 2134: 2095: 2091: 2060:(10): 2656–64. 2046: 2042: 2035: 2021: 2017: 2010: 1994: 1990: 1943: 1939: 1894: 1890: 1884: 1883: 1879: 1834: 1830: 1790: 1786: 1739: 1735: 1680: 1676: 1645: 1641: 1610: 1606: 1597: 1595: 1580: 1574: 1570: 1531: 1527: 1482: 1478: 1433: 1426: 1403:(43): 33820–7. 1389: 1385: 1378: 1356: 1352: 1313: 1309: 1296: 1295: 1291: 1284: 1266: 1262: 1251: 1235: 1228: 1183: 1179: 1134: 1130: 1115: 1093: 1089: 1062: 1058: 1048: 1046: 1037: 1036: 1032: 1028: 1016: 1000:protein folding 969:Crohn's disease 945: 913: 855:signal sequence 847: 799:Golgi apparatus 787: 767: 761: 751:Skeletal muscle 744: 731:gluconeogenesis 685:Golgi apparatus 669: 584:Manufacture of 581:-Golgi network. 573:enzymes with a 569:Manufacture of 532:Golgi apparatus 449: 405: 398: 391: 389:Golgi apparatus 381: 371: 360: 359:on the rough ER 349: 342: 335: 325: 312: 294:Keith R. Porter 209:red blood cells 177:protein folding 173:eukaryotic cell 141: 87:Golgi apparatus 43: 34: 28: 23: 22: 15: 12: 11: 5: 3326: 3316: 3315: 3298: 3297: 3295: 3294: 3289: 3283: 3281: 3277: 3276: 3274: 3273: 3268: 3263: 3262: 3261: 3256: 3246: 3245: 3244: 3239: 3234: 3224: 3218: 3216: 3215:Other internal 3212: 3211: 3209: 3208: 3203: 3202: 3201: 3196: 3191: 3186: 3181: 3176: 3171: 3166: 3161: 3151: 3145: 3143: 3137: 3136: 3134: 3133: 3132: 3131: 3126: 3116: 3115: 3114: 3109: 3104: 3099: 3089: 3084: 3083: 3082: 3077: 3072: 3067: 3057: 3052: 3047: 3042: 3036: 3034: 3028: 3027: 3025: 3024: 3023: 3022: 3017: 3012: 3007: 3002: 2992: 2991: 2990: 2985: 2980: 2975: 2970: 2965: 2955: 2950: 2945: 2940: 2935: 2930: 2924: 2922: 2914: 2913: 2901: 2900: 2893: 2886: 2878: 2872: 2871: 2859: 2849: 2848: 2843: 2836: 2835:External links 2833: 2831: 2830: 2786: 2771: 2722: 2673: 2644:(3): e154068. 2624: 2575: 2546:(8): 455–467. 2526: 2467: 2416: 2358: 2337:(6): 3983–90. 2317: 2268: 2239:(3): 889–902. 2219: 2184: 2132: 2089: 2040: 2034:978-0321909077 2033: 2015: 2008: 1988: 1937: 1888: 1877: 1828: 1801:(2): 123–134. 1784: 1753:(3): 553–562. 1733: 1674: 1639: 1604: 1568: 1541:(18): 188101. 1525: 1476: 1424: 1383: 1376: 1350: 1323:(3): 489–503. 1307: 1289: 1282: 1260: 1249: 1226: 1177: 1128: 1113: 1087: 1056: 1029: 1027: 1024: 1023: 1022: 1015: 1012: 944: 941: 912: 909: 846: 843: 813:family member 786: 783: 757:Main article: 743: 740: 668: 665: 664: 663: 662: 661: 615: 597: 594:signal peptide 582: 528:parking garage 501:signal peptide 471:; also called 448: 445: 311: 308: 267:detoxification 143: 142: 140: 139: 134: 129: 124: 110: 105: 100: 95: 90: 84: 79: 74: 68: 63: 56: 48: 47: 39: 38: 26: 9: 6: 4: 3: 2: 3325: 3314: 3311: 3310: 3308: 3293: 3290: 3288: 3285: 3284: 3282: 3278: 3272: 3269: 3267: 3264: 3260: 3257: 3255: 3252: 3251: 3250: 3247: 3243: 3240: 3238: 3235: 3233: 3230: 3229: 3228: 3225: 3223: 3220: 3219: 3217: 3213: 3207: 3204: 3200: 3197: 3195: 3192: 3190: 3189:Proteinoplast 3187: 3185: 3182: 3180: 3177: 3175: 3172: 3170: 3167: 3165: 3162: 3160: 3157: 3156: 3155: 3152: 3150: 3149:Mitochondrion 3147: 3146: 3144: 3142: 3141:Endosymbionts 3138: 3130: 3127: 3125: 3124:Lamellipodium 3122: 3121: 3120: 3117: 3113: 3110: 3108: 3105: 3103: 3100: 3098: 3095: 3094: 3093: 3090: 3088: 3085: 3081: 3078: 3076: 3073: 3071: 3068: 3066: 3063: 3062: 3061: 3058: 3056: 3053: 3051: 3048: 3046: 3043: 3041: 3040:Microfilament 3038: 3037: 3035: 3033: 3029: 3021: 3018: 3016: 3013: 3011: 3008: 3006: 3003: 3001: 2998: 2997: 2996: 2993: 2989: 2986: 2984: 2981: 2979: 2976: 2974: 2971: 2969: 2966: 2964: 2961: 2960: 2959: 2956: 2954: 2953:Autophagosome 2951: 2949: 2946: 2944: 2941: 2939: 2936: 2934: 2931: 2929: 2928:Cell membrane 2926: 2925: 2923: 2921: 2918:Endomembrane 2915: 2911: 2907: 2899: 2894: 2892: 2887: 2885: 2880: 2879: 2876: 2870: 2866: 2863: 2860: 2858: 2854: 2851: 2850: 2847: 2844: 2842: 2839: 2838: 2826: 2822: 2818: 2814: 2810: 2806: 2802: 2798: 2790: 2782: 2775: 2767: 2763: 2758: 2753: 2749: 2745: 2742:(5): 743–56. 2741: 2737: 2733: 2726: 2718: 2714: 2709: 2704: 2700: 2696: 2692: 2688: 2684: 2677: 2669: 2665: 2660: 2655: 2651: 2647: 2643: 2639: 2635: 2628: 2620: 2616: 2611: 2606: 2602: 2598: 2594: 2590: 2586: 2579: 2571: 2567: 2562: 2557: 2553: 2549: 2545: 2541: 2537: 2530: 2522: 2518: 2513: 2508: 2504: 2500: 2495: 2490: 2486: 2482: 2478: 2471: 2463: 2459: 2454: 2449: 2444: 2439: 2435: 2431: 2427: 2420: 2412: 2408: 2403: 2398: 2394: 2390: 2386: 2382: 2378: 2374: 2370: 2362: 2354: 2350: 2345: 2340: 2336: 2332: 2328: 2321: 2313: 2309: 2304: 2299: 2295: 2291: 2287: 2283: 2279: 2272: 2264: 2260: 2255: 2250: 2246: 2242: 2238: 2234: 2230: 2223: 2215: 2211: 2207: 2203: 2199: 2195: 2188: 2180: 2176: 2172: 2168: 2164: 2160: 2156: 2152: 2148: 2144: 2136: 2128: 2124: 2120: 2116: 2112: 2108: 2104: 2100: 2093: 2085: 2081: 2076: 2071: 2067: 2063: 2059: 2055: 2051: 2044: 2036: 2030: 2026: 2019: 2011: 2009:9780080919317 2005: 2001: 2000: 1992: 1984: 1980: 1976: 1972: 1968: 1964: 1960: 1956: 1952: 1948: 1941: 1933: 1929: 1924: 1919: 1915: 1911: 1907: 1903: 1899: 1892: 1881: 1873: 1869: 1864: 1859: 1855: 1851: 1847: 1843: 1839: 1832: 1824: 1820: 1816: 1812: 1808: 1804: 1800: 1796: 1788: 1780: 1776: 1771: 1766: 1761: 1756: 1752: 1748: 1744: 1737: 1729: 1725: 1720: 1715: 1711: 1707: 1702: 1697: 1693: 1689: 1685: 1678: 1670: 1666: 1662: 1658: 1654: 1650: 1643: 1635: 1631: 1627: 1623: 1620:(9): 483–90. 1619: 1615: 1608: 1593: 1589: 1588: 1583: 1578: 1572: 1564: 1560: 1556: 1552: 1548: 1544: 1540: 1536: 1529: 1521: 1517: 1512: 1507: 1503: 1499: 1496:(2): 285–96. 1495: 1491: 1487: 1480: 1472: 1468: 1464: 1460: 1455: 1450: 1446: 1442: 1438: 1431: 1429: 1420: 1416: 1411: 1406: 1402: 1398: 1394: 1387: 1379: 1373: 1369: 1364: 1363: 1354: 1346: 1342: 1338: 1334: 1330: 1326: 1322: 1318: 1311: 1303: 1299: 1293: 1285: 1279: 1274: 1273: 1264: 1256: 1252: 1246: 1242: 1241: 1233: 1231: 1222: 1218: 1213: 1208: 1204: 1200: 1197:(5): 727–50. 1196: 1192: 1188: 1181: 1173: 1169: 1164: 1159: 1155: 1151: 1148:(3): 233–46. 1147: 1143: 1139: 1132: 1124: 1120: 1116: 1110: 1106: 1102: 1098: 1091: 1083: 1079: 1075: 1071: 1067: 1060: 1044: 1040: 1034: 1030: 1021: 1018: 1017: 1011: 1009: 1005: 1001: 997: 993: 989: 985: 981: 977: 972: 970: 966: 962: 958: 954: 949: 940: 938: 934: 930: 925: 917: 908: 906: 902: 898: 893: 889: 885: 879: 876: 872: 868: 864: 860: 856: 852: 851:glycoproteins 842: 840: 836: 832: 828: 824: 820: 816: 812: 808: 804: 800: 796: 792: 782: 780: 776: 772: 766: 760: 752: 748: 739: 736: 732: 728: 724: 719: 715: 710: 706: 702: 698: 694: 693:phospholipids 690: 686: 682: 673: 659: 655: 651: 647: 643: 639: 635: 631: 627: 623: 622: 620: 619:glycosylation 616: 613: 609: 605: 601: 598: 595: 591: 587: 583: 580: 576: 572: 568: 567: 566: 563: 561: 557: 553: 552:Golgi complex 549: 545: 541: 537: 533: 529: 525: 520: 518: 514: 510: 506: 502: 498: 494: 490: 486: 482: 478: 474: 470: 466: 458: 453: 444: 442: 438: 434: 430: 426: 417: 410: 403: 396: 390: 386: 380: 376: 369: 365: 358: 354: 347: 340: 334: 330: 324: 320: 316: 307: 305: 304: 299: 298:Albert Claude 295: 291: 287: 283: 278: 276: 272: 268: 264: 260: 256: 252: 248: 244: 240: 235: 233: 229: 225: 221: 216: 214: 210: 206: 202: 198: 194: 190: 186: 182: 178: 174: 170: 166: 158: 154: 149: 138: 137:Cell membrane 135: 133: 130: 128: 125: 122: 118: 114: 111: 109: 106: 104: 103:Mitochondrion 101: 99: 96: 94: 91: 88: 85: 83: 80: 78: 75: 72: 69: 67: 64: 62: 59: 58: 54: 49: 46: 40: 37: 31: 19: 3169:Gerontoplast 3119:Pseudopodium 3112:Radial spoke 3092:Undulipodium 3032:Cytoskeleton 2948:Parenthesome 2937: 2857:OPM database 2800: 2796: 2789: 2774: 2739: 2735: 2725: 2690: 2686: 2676: 2641: 2637: 2627: 2592: 2588: 2578: 2543: 2539: 2529: 2484: 2480: 2470: 2433: 2429: 2419: 2376: 2372: 2361: 2334: 2330: 2320: 2285: 2281: 2271: 2236: 2232: 2222: 2197: 2193: 2187: 2146: 2142: 2135: 2102: 2098: 2092: 2057: 2053: 2043: 2024: 2018: 1998: 1991: 1950: 1946: 1940: 1908:(7): 891–8. 1905: 1901: 1891: 1880: 1845: 1841: 1831: 1798: 1794: 1787: 1750: 1746: 1736: 1694:(1): 16–27. 1691: 1687: 1677: 1655:(4): 371–8. 1652: 1648: 1642: 1617: 1613: 1607: 1596:. Retrieved 1585: 1571: 1538: 1534: 1528: 1493: 1489: 1479: 1447:(3): 435–9. 1444: 1440: 1400: 1396: 1386: 1361: 1353: 1320: 1316: 1310: 1301: 1292: 1271: 1263: 1239: 1194: 1190: 1180: 1145: 1141: 1131: 1096: 1090: 1073: 1069: 1059: 1047:. Retrieved 1033: 998:involved in 973: 950: 946: 936: 932: 928: 926: 922: 880: 875:cytoskeleton 848: 823:calreticulin 788: 775:muscle cells 770: 768: 680: 678: 645: 578: 564: 521: 472: 468: 464: 462: 429:cytoskeleton 422: 408: 406:   401: 399:   394: 392:   384: 382:   374: 372:   363: 361:   352: 350:   345: 343:   338: 336:   333:Nuclear pore 328: 326:   318: 301: 286:ergastoplasm 285: 279: 236: 217: 196: 192: 188: 184: 168: 164: 162: 157:mitochondria 93:Cytoskeleton 35:Cell biology 3271:Magnetosome 3237:Spliceosome 3164:Chromoplast 3159:Chloroplast 3050:Microtubule 2783:. iBiology. 2379:(1): 3489. 1587:Answers.com 1298:"reticulum" 1076:: 278–289. 1049:21 November 992:translation 978:(UPR) is a 863:amino acids 859:polypeptide 718:calcium ion 497:amino acids 485:translating 292:in 1945 by 251:hepatocytes 237:The outer ( 228:cholesterol 213:spermatozoa 3266:Proteasome 3259:Inclusions 3206:Nitroplast 3199:Apicoplast 3184:Elaioplast 3179:Amyloplast 3174:Leucoplast 3129:Filopodium 3075:Basal body 3065:Centrosome 3015:Peroxisome 3010:Glyoxysome 3000:Melanosome 2910:organelles 2487:: e49682. 1598:2006-09-13 1026:References 996:chaperones 963:. In the 867:translocon 763:See also: 509:translocon 481:translocon 259:metabolism 132:Centrosome 117:organelles 3287:Cell wall 3249:Cytoplasm 3222:Nucleolus 3194:Tannosome 3102:Flagellum 3087:Myofibril 3070:Centriole 3005:Microbody 2978:Phagosome 2503:2050-084X 1710:1535-9476 1082:493441682 815:BiP/Grp78 803:chaperone 791:cisternae 785:Functions 571:lysosomal 517:peptidase 477:ribosomes 425:cisternae 310:Structure 303:reticulum 243:ribosomes 239:cytosolic 201:cisternae 181:organelle 121:cytoplasm 61:Nucleolus 3307:Category 3280:External 3232:Ribosome 2988:Acrosome 2973:Endosome 2968:Lysosome 2865:Archived 2825:25570626 2817:24107777 2766:18775308 2717:21389082 2668:34905513 2619:32651241 2589:Diabetes 2570:34163039 2521:31498082 2462:31498082 2411:30154480 2312:18086916 2263:12631711 2179:22517395 2171:15486293 2127:25858120 2119:22510960 2084:16200199 1975:10864315 1932:12370264 1872:29290583 1815:25355612 1779:28119445 1728:30237200 1669:16806880 1634:15350976 1592:Archived 1563:25396396 1520:23870120 1471:16107069 1463:16901774 1419:10931837 1345:19078317 1255:Archived 1221:13052830 1172:19871454 1123:14283576 1043:Archived 1014:See also 984:proteins 871:vesicles 819:calnexin 795:vesicles 697:steroids 658:nitrogen 652:, and 3- 617:Initial 590:clathrin 586:secreted 469:rough ER 368:Proteins 357:Ribosome 220:proteins 127:Lysosome 71:Ribosome 3254:Cytosol 3154:Plastid 3107:Axoneme 2983:Vacuole 2963:Exosome 2958:Vesicle 2933:Nucleus 2757:2586148 2708:3090193 2659:8803344 2610:7506835 2561:8765009 2512:6763289 2453:6763289 2402:6113206 2381:Bibcode 2353:1740446 2303:2140024 2214:1363623 2151:Bibcode 2143:Science 2075:1236697 1983:4316039 1955:Bibcode 1863:5975648 1795:Traffic 1770:5335585 1719:6317476 1543:Bibcode 1511:3767119 1368:659–666 1337:1423609 1212:2136295 1163:2135493 986:in the 937:CaATiER 933:CaATiER 797:to the 705:ovaries 660:of Asn. 654:glucose 650:mannose 513:nascent 441:cytosol 379:vesicle 323:Nucleus 277:cells. 191:), and 153:nucleus 113:Cytosol 108:Vacuole 77:Vesicle 66:Nucleus 3097:Cilium 2920:system 2823:  2815:  2764:  2754:  2715:  2705:  2666:  2656:  2617:  2607:  2568:  2558:  2519:  2509:  2501:  2460:  2450:  2409:  2399:  2351:  2310:  2300:  2261:  2254:151567 2251:  2212:  2177:  2169:  2125:  2117:  2082:  2072:  2031:  2006:  1981:  1973:  1947:Nature 1930:  1923:151159 1920:  1870:  1860:  1821:  1813:  1777:  1767:  1726:  1716:  1708:  1667:  1632:  1561:  1518:  1508:  1469:  1461:  1417:  1374:  1343:  1335:  1280:  1247:  1219:  1209:  1170:  1160:  1121:  1111:  1080:  707:, and 701:testes 695:, and 689:lipids 431:. 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Index

Smooth endoplasmic reticulum
Cell biology
Animal cell diagram

Nucleolus
Nucleus
Ribosome
Vesicle
Rough endoplasmic reticulum
Golgi apparatus
Cytoskeleton
Smooth endoplasmic reticulum
Mitochondrion
Vacuole
Cytosol
organelles
cytoplasm
Lysosome
Centrosome
Cell membrane

nucleus
mitochondria
eukaryotic cell
protein folding
organelle
cisternae
nuclear membrane
red blood cells
spermatozoa

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