小器官、オルガネラ、細胞小器官、細胞器官

関: cell organ、organellar

WordNet

a specialized part of a cell; analogous to an organ; "the first organelle to be identified was the nucleus" (同)cell organelle, cell_organ

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2016/03/07 10:28:10」(JST)

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In cell biology, an organelle /ɔːrɡəˈnɛl/ is a specialized subunit within a cell that has a specific function. Individual organelles are usually separately enclosed within their own lipid bilayers.

The name organelle comes from the idea that these structures are to cells what an organ is to the body (hence the name organelle, the suffix -elle being a diminutive). Organelles are identified by microscopy, and can also be purified by cell fractionation. There are many types of organelles, particularly in eukaryotic cells. While prokaryotes do not possess organelles per se, some do contain protein-based microcompartments, which are thought to act as primitive organelles.^[1]

History and terminology

In biology organs are defined as confined functional units within an organism.^[2] The analogy of bodily organs to microscopic cellular substructures is obvious, as from even early works, authors of respective textbooks rarely elaborate on the distinction between the two.

Credited as the first^[3]^[4]^[5] to use a diminutive of organ (i.e., little organ) for cellular structures was German zoologist Karl August Möbius (1884), who used the term organula (plural of organulum, the diminutive of Latin organum).^[6] In a footnote, which was published as a correction in the next issue of the journal, he justified his suggestion to call organs of unicellular organisms "organella" since they are only differently formed parts of one cell, in contrast to multicellular organs of multicellular organisms.^{[citation needed]}

Types of organelles

While most cell biologists consider the term organelle to be synonymous with "cell compartment", other cell biologists choose to limit the term organelle to include only those that are DNA-containing, having originated from formerly autonomous microscopic organisms acquired via endosymbiosis.^[7]^[8]^[9]

Under this definition, there would only be two broad classes of organelles (i.e. those that contain their own DNA, and have originated from endosymbiotic bacteria):

mitochondria (in almost all eukaryotes)
plastids^[10] (e.g. in plants, algae, and some protists).

Other organelles are also suggested to have endosymbiotic origins, but do not contain their own DNA (notably the flagellum – see evolution of flagella).

Under the more restricted definition of membrane-bound structures, some parts of the cell do not qualify as organelles. Nevertheless, the use of organelle to refer to non-membrane bound structures such as ribosomes is common.^[11] This has led some texts to delineate between membrane-bound and non-membrane bound organelles.^[12] The non-membrane bound organelles, also called large biomolecular complexes, are large assemblies of macromolecules that carry out particular and specialized functions, but they lack membrane boundaries. Such cell structures include:

large RNA and protein complexes: ribosome, spliceosome, vault
large protein complexes: proteasome, DNA polymerase III holoenzyme, RNA polymerase II holoenzyme, symmetric viral capsids, complex of GroEL and GroES; membrane protein complexes: photosystem I, ATP synthase
large DNA and protein complexes: nucleosome
centriole and microtubule-organizing center (MTOC)
cytoskeleton
flagellum
cellular structure which is not membrane bound and does not have a well-defined structure: nucleolus

Eukaryotic organelles

Eukaryotic cells are structurally complex, and by definition are organized, in part, by interior compartments that are themselves enclosed by lipid membranes that resemble the outermost cell membrane. The larger organelles, such as the nucleus and vacuoles, are easily visible with the light microscope. They were among the first biological discoveries made after the invention of the microscope.

Not all eukaryotic cells have each of the organelles listed below. Exceptional organisms have cells that do not include some organelles that might otherwise be considered universal to eukaryotes (such as mitochondria).^[13] There are also occasional exceptions to the number of membranes surrounding organelles, listed in the tables below (e.g., some that are listed as double-membrane are sometimes found with single or triple membranes). In addition, the number of individual organelles of each type found in a given cell varies depending upon the function of that cell.

**Major eukaryotic organelles**
Organelle	Main function	Structure	Organisms	Notes
chloroplast (plastid)	photosynthesis, traps energy from sunlight	double-membrane compartment	plants, protists (rare kleptoplastic organisms)	has own DNA; theorized to be engulfed by the ancestral eukaryotic cell (endosymbiosis)
endoplasmic reticulum	translation and folding of new proteins (rough endoplasmic reticulum), expression of lipids (smooth endoplasmic reticulum)	single-membrane compartment	all eukaryotes	rough endoplasmic reticulum is covered with ribosomes, has folds that are flat sacs; smooth endoplasmic reticulum has folds that are tubular
Flagellum	locomotion, sensory		some eukaryotes
Golgi apparatus	sorting, packaging, processing and modification of proteins	single-membrane compartment	all eukaryotes	cis-face (convex) nearest to rough endoplasmic reticulum; trans-face (concave) farthest from rough endoplasmic reticulum
mitochondria	energy production from the oxidation of glucose substances and the release of adenosine triphosphate	double-membrane compartment	most eukaryotes	has own DNA; theorized to be engulfed by an ancestral eukaryotic cell (endosymbiosis)
vacuole	storage, transportation, helps maintain homeostasis	single-membrane compartment	eukaryotes
nucleus	DNA maintenance, controls all activities of the cell, RNA transcription	double-membrane compartment	all eukaryotes	contains bulk of genome

Mitochondria and chloroplasts, which have double-membranes and their own DNA, are believed to have originated from incompletely consumed or invading prokaryotic organisms, which were adopted as a part of the invaded cell. This idea is supported in the Endosymbiotic theory.

**Minor eukaryotic organelles and cell components**
Organelle/Macromolecule	Main function	Structure	Organisms
acrosome	helps spermatozoa fuse with ovum	single-membrane compartment	many animals
autophagosome	vesicle that sequesters cytoplasmic material and organelles for degradation	double-membrane compartment	all eukaryotes
centriole	anchor for cytoskeleton, organizes cell division by forming spindle fibers	Microtubule protein	animals
cilium	movement in or of external medium; "critical developmental signaling pathway".^[14]	Microtubule protein	animals, protists, few plants
eyespot apparatus	detects light, allowing phototaxis to take place		green algae and other unicellular photosynthetic organisms such as euglenids
glycosome	carries out glycolysis	single-membrane compartment	Some protozoa, such as Trypanosomes.
glyoxysome	conversion of fat into sugars	single-membrane compartment	plants
hydrogenosome	energy & hydrogen production	double-membrane compartment	a few unicellular eukaryotes
lysosome	breakdown of large molecules (e.g., proteins + polysaccharides)	single-membrane compartment	most eukaryotes
melanosome	pigment storage	single-membrane compartment	animals
mitosome	probably plays a role in Iron-sulfur cluster (Fe-S) assembly	double-membrane compartment	a few unicellular eukaryotes that lack mitochondria
myofibril	myocyte contraction	bundled filaments	animals
nematocyst	stinging	coiled hollow tubule	Cnidarians
nucleolus	pre-ribosome production	protein-DNA-RNA	most eukaryotes
parenthesome	not characterized	not characterized	fungi
peroxisome	breakdown of metabolic hydrogen peroxide	single-membrane compartment	all eukaryotes
proteasome	degradation of unneeded or damaged proteins by proteolysis	very large protein complex	All eukaryotes, all archaea, some bacteria
ribosome (80S)	translation of RNA into proteins	RNA-protein	all eukaryotes
vesicle	material transport	single-membrane compartment	all eukaryotes

Other related structures:

cytosol
endomembrane system
nucleosome
microtubule
cell membrane

(A) Electron micrograph of Halothiobacillus neapolitanus cells, arrows highlight carboxysomes. (B) Image of intact carboxysomes isolated from H. neapolitanus. Scale bars are 100 nm.^[15]

Prokaryotic organelles

Prokaryotes are not as structurally complex as eukaryotes, and were once thought not to have any internal structures enclosed by lipid membranes. In the past, they were often viewed as having little internal organization, but slowly, details are emerging about prokaryotic internal structures. An early false turn was the idea developed in the 1970s that bacteria might contain membrane folds termed mesosomes, but these were later shown to be artifacts produced by the chemicals used to prepare the cells for electron microscopy.^[16]

However, more recent research has revealed that at least some prokaryotes have microcompartments such as carboxysomes. These subcellular compartments are 100–200 nm in diameter and are enclosed by a shell of proteins.^[1] Even more striking is the description of membrane-bound magnetosomes in bacteria, reported in 2006,^[17]^[18] as well as the nucleus-like structures of the Planctomycetes that are surrounded by lipid membranes, reported in 2005.^[19]

**Prokaryotic organelles and cell components**
Organelle/Macromolecule	Main function	Structure	Organisms
carboxysome	carbon fixation	protein-shell compartment	some bacteria
chlorosome	photosynthesis	light harvesting complex	green sulfur bacteria
flagellum	movement in external medium	protein filament	some prokaryotes and eukaryotes
magnetosome	magnetic orientation	inorganic crystal, lipid membrane	magnetotactic bacteria
nucleoid	DNA maintenance, transcription to RNA	DNA-protein	prokaryotes
plasmid	DNA exchange	circular DNA	some bacteria
ribosome (70S)	translation of RNA into proteins	RNA-protein	bacteria and archaea
thylakoid	photosynthesis	photosystem proteins and pigments	mostly cyanobacteria
mesosomes	functions of Golgi bodies, centrioles, etc.	small irregular shaped organelle containing ribosomes	present in most prokaryotic cells
Pilus	Adhesion to other cells for conjugation or to a solid substrate to create motile forces.	a hair-like appendage sticking out (though partially embedded into) the plasma membrane	prokaryotic cells

Proteins and organelles

The function of a protein is closely correlated with the organelle in which it resides. Some methods were proposed for predicting the organelle in which an uncharacterized protein is located according to its amino acid composition^[20]^[21] and some methods were based on pseudo amino acid composition.^[22]^[23]^[24]^[25]

References

^ ^a ^b Kerfeld, C. A.; Sawaya, M. R; Tanaka, S; Nguyen, C. V.; Phillips, M; Beeby, M; Yeates, T. O. (5 August 2005). "Protein structures forming the shell of primitive bacterial organelles.". Science 309 (5736): 936–8. Bibcode:2005Sci...309..936K. doi:10.1126/science.1113397. PMID 16081736.
^ Lynsey Peterson (2010-04-17). "Mastering the Parts of a Cell". Lesson Planet. Retrieved 2010-04-19.
^ Bütschli, O. (1888). Dr. H. G. Bronn's Klassen u. Ordnungen des Thier-Reichs wissenschaftlich dargestellt in Wort und Bild. Erster Band. Protozoa. Dritte Abtheilung: Infusoria und System der Radiolaria. p. 1412. Die Vacuolen sind demnach in strengem Sinne keine beständigen Organe oder O r g a n u l a (wie Möbius die Organe der Einzelligen im Gegensatz zu denen der Vielzelligen zu nennen vorschlug).
^ Amer. Naturalist. 23, 1889, p. 183: "It may possibly be of advantage to use the word organula here instead of organ, following a suggestion by Möbius. Functionally differentiated multicellular aggregates in multicellular forms or metazoa are in this sense organs, while, for functionally differentiated portions of unicellular organisms or for such differentiated portions of the unicellular germ-elements of metazoa, the diminutive organula is appropriate." Cited after: Oxford English Dictionary online, entry for "organelle".
^ 'Journal de l'anatomie et de la physiologie normales et pathologiques de l'homme et des animaux' at Google Books
^ Möbius, K. (September 1884). "Das Sterben der einzelligen und der vielzelligen Tiere. Vergleichend betrachtet". Biologisches Centralblatt 4 (13, 14): 389–392, 448. Während die Fortpflanzungszellen der vielzelligen Tiere unthätig fortleben bis sie sich loslösen, wandern und entwickeln, treten die einzelligen Tiere auch durch die an der Fortpflanzung beteiligten Leibesmasse in Verkehr mit der Außenwelt und viele bilden sich dafür auch besondere Organula". Footnote on p. 448: "Die Organe der Heteroplastiden bestehen aus vereinigten Zellen. Da die Organe der Monoplastiden nur verschieden ausgebildete Teile e i n e r Zelle sind schlage ich vor, sie „Organula“ zu nennen
^ Keeling, Pj; Archibald, Jm (2008). "Organelle evolution: what's in a name?". Current Biology 18 (8): R345–7. doi:10.1016/j.cub.2008.02.065. PMID 18430636.
^ Imanian B, Carpenter KJ, Keeling PJ (2007). "Mitochondrial genome of a tertiary endosymbiont retains genes for electron transport proteins". The Journal of eukaryotic microbiology 54 (2): 146–53. doi:10.1111/j.1550-7408.2007.00245.x. PMID 17403155.
^ Mullins, Christopher (2004). "Theory of Organelle Biogenesis: A Historical Perspective". The Biogenesis of Cellular Organelles. Springer Science+Business Media, National Institutes of Health. ISBN 0-306-47990-7.
^ C.Michael Hogan. 2010. Deoxyribonucleic acid. Encyclopedia of Earth. National Council for Science and the Environment. S. Draggan and C. Cleveland (eds.). Washington DC
^ Campbell and Reece, Biology 6th edition, Benjamin Cummings, 2002
^ Cormack, David H. (1984) Introduction to Histology, Lippincott, ISBN 0397521146
^ Fahey RC, Newton GL, Arrack B, Overdank-Bogart T, Baley S (1984). "Entamoeba histolytica: a eukaryote without glutathione metabolism". Science 224 (4644): 70–72. Bibcode:1984Sci...224...70F. doi:10.1126/science.6322306. PMID 6322306.
^ Badano, Jose L.; Norimasa Mitsuma; Phil L. Beales; Nicholas Katsanis (September 2006). "The Ciliopathies: An Emerging Class of Human Genetic Disorders". Annual Review of Genomics and Human Genetics 7: 125–148. doi:10.1146/annurev.genom.7.080505.115610. PMID 16722803.
^ Tsai Y, Sawaya MR, Cannon GC, Cai F, Williams EB, Heinhorst S, Kerfeld CA, Yeates TO (2007). "Structural Analysis of CsoS1A and the Protein Shell of the Halothiobacillus neapolitanus Carboxysome". PLoS Biology 5 (6): e144. doi:10.1371/journal.pbio.0050144. PMC 1872035. PMID 17518518.
^ Ryter A (1988). "Contribution of new cryomethods to a better knowledge of bacterial anatomy". Ann. Inst. Pasteur Microbiol. 139 (1): 33–44. doi:10.1016/0769-2609(88)90095-6. PMID 3289587.
^ Komeili A, Li Z, Newman DK, Jensen GJ (2006). "Magnetosomes are cell membrane invaginations organized by the actin-like protein MamK". Science 311 (5758): 242–5. Bibcode:2006Sci...311..242K. doi:10.1126/science.1123231. PMID 16373532.
^ Scheffel A, Gruska M, Faivre D, Linaroudis A, Plitzko JM, Schüler D (2006). "An acidic protein aligns magnetosomes along a filamentous structure in magnetotactic bacteria". Nature 440 (7080): 110–4. Bibcode:2006Natur.440..110S. doi:10.1038/nature04382. PMID 16299495.
^ Fuerst JA (2005). "Intracellular compartmentation in planctomycetes". Annu. Rev. Microbiol. 59: 299–328. doi:10.1146/annurev.micro.59.030804.121258. PMID 15910279.
^ Cedano, J.; Aloy, P.; P'erez-Pons, J. A.; Querol, E. (1997). "Relation between amino acid composition and cellular location of proteins". J. Mol. Biol. 266 (3): 594–600. doi:10.1006/jmbi.1996.0804. PMID 9067612.
^ Chou, K. C.; Elrod, D. W. (1999). "Protein subcellular location prediction". Protein Engineering 12 (2): 107–118. doi:10.1093/protein/12.2.107. PMID 10195282.
^ Chou, KC (2001). "Prediction of protein cellular attributes using pseudo-amino acid composition". Proteins 43 (3): 246–55. doi:10.1002/prot.1035. PMID 11288174.
^ Mundra, P.; Kumar, M.; Kumar, K. K.; Jayaraman, V. K.; Kulkarni, B. D. (2007). "Using pseudo amino acid composition to predict protein subnuclear localization: Approached with PSSM". Pattern Recognition Letters 28 (13): 1610–1615. doi:10.1016/j.patrec.2007.04.001.
^ Du, P.; Cao, S.; Li, Y. (2009). "SubChlo: predicting protein subchloroplast locations with pseudo-amino acid composition and the evidence-theoretic K-nearest neighbor (ET-KNN) algorithm". Journal of Theoretical Biology 261 (2): 330–335. doi:10.1016/j.jtbi.2009.08.004.
^ Li, F. M.; Li, Q. Z. (2008). "Predicting protein subcellular location using Chou's pseudo amino acid composition and improved hybrid approach". Protein & Peptide Letters 15 (6): 612–616. doi:10.2174/092986608784966930. PMID 18680458.

External links

Tree of Life project: Eukaryotes

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リンク元	「細胞小器官」「小器官」「organellar」「オルガネラ」「cell organ」
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	Library resources about; Organelle
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Cell biology
	The animal cell
	Components of a typical animal cell: Nucleolus; Nucleus; Ribosome (little dots); Vesicle; Rough endoplasmic reticulum; Golgi apparatus (or "Golgi body"); Cytoskeleton; Smooth endoplasmic reticulum; Mitochondrion; Vacuole; Cytosol (fluid that contains organelles); Lysosome; Centrosome; Cell membrane;