actin-related protein 2-3 complex

アクチン関連タンパク質 2/3複合体

WordNet

1. something that people do or cause to happen (同)deed, human action, human_activity
2. behave in a certain manner; show a certain behavior; conduct or comport oneself; "You should act like an adult"; "Dont behave like a fool"; "What makes her do this way?"; "The dog acts ferocious, but he is really afraid of people" (同)behave, do
3. pretend to have certain qualities or state of mind; "He acted the idiot"; "She plays deaf when the news are bad" (同)play, act_as
4. a legal document codifying the result of deliberations of a committee or society or legislative body (同)enactment
5. a short theatrical performance that is part of a longer program; "he did his act three times every evening"; "she had a catchy little routine"; "it was one of the best numbers he ever did" (同)routine, number, turn, bit
6. a manifestation of insincerity; "he put on quite an act for her benefit"
7. a subdivision of a play or opera or ballet
8. discharge ones duties; "She acts as the chair"; "In what capacity are you acting?"
9. perform an action, or work out or perform (an action); "think before you act"; "We must move quickly"; "The governor should act on the new energy bill"; "The nanny acted quickly by grabbing the toddler and covering him with a wet towel" (同)move
10. play a role or part; "Gielgud played Hamlet"; "She wants to act Lady Macbeth, but she is too young for the role"; "She played the servant to her husbands master" (同)play, represent
11. perform on a stage or theater; "She acts in this play"; "He acted in `Julius Caesar"; "I played in `A Christmas Carol" (同)play, roleplay, playact
12. be engaged in an activity, often for no particular purpose other than pleasure
13. be suitable for theatrical performance; "This scene acts well"
14. complicated in structure; consisting of interconnected parts; "a complex set of variations based on a simple folk melody"; "a complex mass of diverse laws and customs"
15. a compound described in terms of the central atom to which other atoms are bound or coordinated (同)coordination_compound
16. a conceptual whole made up of complicated and related parts; "the complex of shopping malls, houses, and roads created a new town" (同)composite
17. (psychoanalysis) a combination of emotions and impulses that have been rejected from awareness but still influence a persons behavior
18. have or establish a relationship to; "She relates well to her peers"
19. be in a relationship with; "How are these two observations related?" (同)interrelate
20. give an account of; "The witness related the events"
21. connected by kinship, common origin, or marriage
22. being connected either logically or causally or by shared characteristics ; "painting and the related arts"; "school-related activities"; "related to micelle formation is the...ability of detergent actives to congregate at oil-water interfaces" (同)related to
23. any of a large group of nitrogenous organic compounds that are essential constituents of living cells; consist of polymers of amino acids; essential in the diet of animals for growth and for repair of tissues; can be obtained from meat and eggs and milk and legumes; "a diet high in protein"

PrepTutorEJDIC

1. 『行い』,『行為』 / 《しばしばA-》法令,条令 / 《しばしばA-》(芝居の1つの)『幕』;(演芸などの)出し物の一つ / (見せかけの)お芝居,演技 / 『行動する』,行う / 《様態を表す副詞[句]を伴って》わざとふるまう,見せかける / 〈人・物が〉(…として)動く,役割をはたす《+『as』+『名』》 / 〈薬が〉『作用する』,きく / 舞台に立つ,出演する / 〈…の役〉'を'『演ずる』;〈劇〉'を'上演する / …‘に'ふさわしくふるまう / …‘に'見せかける,‘の'ふりをする
2. 『いくつかの部分から成る』,複合の,合成の / 『複雑な』,入りくんだ,こみいった(complicated) / 複合体,合成物 / コンプレックス,複合(抑圧されて心に残った複雑なしこり)
3. (…に)〈事件・事情など〉‘を'『物語る』《+『名』+『to』+『名』》 / (…と)…‘を'『関係させる』,結びつける《+『名』+『to(with)』+『名』》 / 〈物・事が〉(…に)関係がある,かかわる《+『to(with)』+『名(wh・節・句)』》 / (…に)〈人が〉なじむ,順応する《+『to』+『名』》
4. 関係のある
5. 蛋白(たんばく)質

Wikipedia preview

出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2012/12/11 06:51:59」(JST)

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[Wiki en表示]

Atomic structure of bovine Arp2/3 complex (PDB code: 1k8k). Color coding for subunits: Arp3, orange; Arp2, marine (subunits 1 & 2 not resolved and thus not shown); p40, green; p34, ice blue; p20, dark blue; p21, magenta; p16, yellow.

Arp2/3 complex is a seven-subunit protein that plays a major role in the regulation of the actin cytoskeleton. It is a major component of the actin cytoskeleton and is found in most in actin cytoskeleton-containing eukaryotic cells. Two of its subunits, the Actin-Related Proteins ARP2 and ARP3 closely resemble the structure of monomeric actin and serve as nucleation sites for new actin filaments. The complex binds to the sides of existing ("mother") filaments and initiates growth of a new ("daughter") filament at a distinctive 70 degree angle from the mother. Branched actin networks are created as a result of this nucleation of new filaments. The regulation of rearrangements of the actin cytoskeleton is important for processes like cell locomotion, phagocytosis, and intracellular motility of lipid vesicles.

The Arp2/3 complex was named after it was identified by affinity chromatography from Acanthamoeba castellanii, though it had been previously isolated in 1989 in a search for proteins that bind to actin filaments in Drosophila melanogaster embryos . It is found in most eukaryotic organisms, but absent from a number of Chromalveolates and plants.

Mechanisms of actin polymerization by Arp2/3

Side branching model of the Arp2/3 complex. Activated Arp2/3 complex binds to the side of a "mother" actin filament. Both Arp2 and Arp3 form the first two subunits in the new "daughter" filament.

Barbed end branching model of the Arp2/3 complex. Activated Arp2/3 competes with capping proteins to bind to the barbed end of an actin filament. Arp2 remains bound to the mother filament, while Arp3 is outside. The two Arp subunits form the first subunits of each branch and the two branches continue to grow by addition of G-actin to each Arp

Many actin-related molecules create a free barbed end for polymerization by uncapping or severing pre-existing filaments and using these as nucleation cores. However, the Arp2/3 complex stimulates actin polymerization by creating a new nucleation core. Actin nucleation is an initial step in the formation of an actin filament. The nucleation core activity of Arp2/3 is activated by members of the Wiskott-Aldrich syndrome family protein (WASP, N-WASP, WAVE, and WASH proteins). The V domain of a WASP protein interacts with actin monomers while the CA region associates with the Arp2/3 complex to create a nucleation core. However, de novo nucleation followed by polymerization is not sufficient to form integrated actin networks, since these newly synthesized polymers would not be associated with pre-existing filaments. Thus, the Arp2/3 complex binds to pre-existing filaments so that the new filaments can grow on the old ones and form a functional actin cytoskeleton. Capping proteins limit actin polymerization to the region activated by the Arp2/3 complex, and the elongated filament ends are recapped to prevent depolymerization and thus conserve the actin filament.

The Arp2/3 complex simultaneously controls nucleation of actin polymerization and branching of filaments. Moreover, autocatalysis is observed during Arp2/3-mediated actin polymerization. In this process, the newly formed filaments activate other Arp2/3 complexes, facilitating the formation of branched filaments.

The mechanisms of actin polymerization by Arp2/3 has been the subject of dispute in the recent years. The question is where the complex binds the filament and how it nucleates a "daughter" filament. Historically two models have been proposed to describe the formation of branched filaments:

Side branching model

In the side branching (or dendritic nucleation) model, the Arp2/3 complex binds to the side of a pre-existing ("mother") filament at a point different from the nucleation site. Arp2/3 thus has two actin-binding sites — one to bind to the pre-existing actin filament and the other for the nucleation of a branched filament. Recent structural data from high-resolution electron microscopy, together with biochemical data on the filament nucleation and capping mechanisms of the Arp2/3 complex, provide strong support for this model.

Barbed end branching model

In the barbed end branching model, Arp2/3 associates at the barbed end of growing filaments, allowing for the elongation of the original filament and the formation of a branched filament, suggesting that branching is induced with Arp2 and Arp3 being incorporated in two different actin filaments. However, this model is only based on kinetic analysis and lacks support from structural data.

Cellular uses of Arp2/3

The Arp2/3 complex appears to be important in a variety of specialized cell functions that involve the actin cytoskeleton. The complex is found in cellular regions characterized by dynamic actin filament activity; in macropinocytic cups, in the leading edge of motile cells (lamellipodia), and in motile actin patches in yeast. In mammals and the social amoeba Dictyostelium discoideum it is required for phagocytosis. The complex has also been shown to be involved in the establishment of cell polarity and the migration of fibroblast monolayers in a wound-healing model. In mammlian oocytes, Arp2/3 complex involved into the oocyte asymmetric division and polar body emission, which are resulted from the failure of spindle migration (a unique feathure of oocyte division) and cytokinesis. Moreover, enteropathogenic organisms like Listeria monocytogenes and Shigella use the Arp2/3 complex for actin-polymerization dependent rocketing movements. The Arp2/3 complex also regulates the intracellular motility of endosomes, lysosomes, pinocytic vesicles and mitochondria. Moreover, recent studies show that the Arp2/3 complex is essential for proper polar cell expansion in plants. Arp2/3 mutations in Arabidopsis thaliana result in abnormal filament organization, which in turn affects the expansion of trichomes, pavement cells, hypocotyl cells, and root hair cells.

References

1. ^a Robinson RC, Turbedsky K, Kaiser DA, et al. (November 2001). "Crystal structure of Arp2/3 complex". Science 294 (5547): 1679–84. doi:10.1126/science.1066333. PMID 11721045. http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=11721045.
2. ^{a b} Suetsugu S, Miki H, Takenawa T (March 2002). "Spatial and temporal regulation of actin polymerization for cytoskeleton formation through Arp2/3 complex and WASP/WAVE proteins". Cell Motil. Cytoskeleton 51 (3): 113–22. doi:10.1002/cm.10020. PMID 11921168.
3. ^a Aguda AH, Burtnick LD, Robinson RC (March 2005). "The state of the filament". EMBO Rep. 6 (3): 220–6. doi:10.1038/sj.embor.7400363. PMC 1299273. PMID 15741975. //www.ncbi.nlm.nih.gov/pmc/articles/PMC1299273/.
4. ^a Egile C, Rouiller I, Xu XP, Volkmann N, Li R, Hanein D (November 2005). "Mechanism of filament nucleation and branch stability revealed by the structure of the Arp2/3 complex at actin branch junctions". PLoS Biol. 3 (11): e383. doi:10.1371/journal.pbio.0030383. PMC 1278936. PMID 16262445. http://dx.plos.org/10.1371/journal.pbio.0030383.
5. ^a Bannigan A, Baskin TI (December 2005). "Directional cell expansion—turning toward actin". Curr. Opin. Plant Biol. 8 (6): 619–24. doi:10.1016/j.pbi.2005.09.002. PMID 16181803. http://linkinghub.elsevier.com/retrieve/pii/S1369-5266(05)00124-X.
6. ^a Mathur J (April 2005). "The ARP2/3 complex: giving plant cells a leading edge". BioEssays 27 (4): 377–87. doi:10.1002/bies.20206. PMID 15770684.
7. ^a Xu J, Scheres B (December 2005). "Cell polarity: ROPing the ends together". Curr. Opin. Plant Biol. 8 (6): 613–8. doi:10.1016/j.pbi.2005.09.003. PMID 16182602. http://linkinghub.elsevier.com/retrieve/pii/S1369-5266(05)00127-5.
8. ^a Warren DT, Andrews PD, Gourlay CW, Ayscough KR (April 2002). "Sla1p couples the yeast endocytic machinery to proteins regulating actin dynamics". J. Cell. Sci. 115 (Pt 8): 1703–15. PMID 11950888. http://jcs.biologists.org/cgi/pmidlookup?view=long&pmid=11950888.
9. ^a May RC, Caron E, Hall A, Machesky LM (April 2000). "Involvement of the Arp2/3 complex in phagocytosis mediated by FcgammaR or CR3". Nat. Cell Biol. 2 (4): 246–8. doi:10.1038/35008673. PMID 10783245.
10. ^a Insall R, Müller-Taubenberger A, Machesky L, et al. (November 2001). "Dynamics of the Dictyostelium Arp2/3 complex in endocytosis, cytokinesis, and chemotaxis". Cell Motil. Cytoskeleton 50 (3): 115–28. doi:10.1002/cm.10005. PMID 11807934.
11. ^a Magdalena J, Millard TH, Etienne-Manneville S, Launay S, Warwick HK, Machesky LM (February 2003). "Involvement of the Arp2/3 complex and Scar2 in Golgi polarity in scratch wound models". Mol. Biol. Cell 14 (2): 670–84. doi:10.1091/mbc.E02-06-0345. PMC 150000. PMID 12589062. http://www.molbiolcell.org/cgi/pmidlookup?view=long&pmid=12589062.
12. ^a Volkmann N, Amann KJ, Stoilova-McPhie S, et al. (September 2001). "Structure of Arp2/3 complex in its activated state and in actin filament branch junctions". Science 293 (5539): 2456–9. doi:10.1126/science.1063025. PMID 11533442. http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=11533442.
13. ^a Dayel MJ, Mullins RD (April 2004). "Activation of Arp2/3 complex: addition of the first subunit of the new filament by a WASP protein triggers rapid ATP hydrolysis on Arp2". PLoS Biol. 2 (4): E91. doi:10.1371/journal.pbio.0020091. PMC 387265. PMID 15094799. http://dx.plos.org/10.1371/journal.pbio.0020091.
14. ^a Machesky LM, Atkinson SJ, Ampe C, Vandekerckhove J, Pollard TD (October 1994). "Purification of a cortical complex containing two unconventional actins from Acanthamoeba by affinity chromatography on profilin-agarose". J. Cell Biol. 127 (1): 107–15. doi:10.1083/jcb.127.1.107. PMC 2120189. PMID 7929556. http://www.jcb.org/cgi/pmidlookup?view=long&pmid=7929556.
15. ^a Miller KG, Field CM, Alberts BM (December 1989). "Actin-binding proteins from Drosophila embryos: a complex network of interacting proteins detected by F-actin affinity chromatography". J. Cell Biol. 109 (6 Pt 1): 2963–75. doi:10.1083/jcb.109.6.2963. PMC 2115944. PMID 2512303. http://www.jcb.org/cgi/pmidlookup?view=long&pmid=2512303.
16. ^a Veltman DM, Insall RH (August 2010). "WASP family proteins: their evolution and its physiological implications". Mol. Biol. Cell 21 (16): 2880–93. doi:10.1091/mbc.E10-04-0372. PMC 2921111. PMID 20573979. http://www.molbiolcell.org/cgi/pmidlookup?view=long&pmid=20573979.

External links

• MBInfo - Arp2/3 Mediated Nucleation
• Arp2-3+Complex at the US National Library of Medicine Medical Subject Headings (MeSH)
• Arp2/3 filament nucleation model, illustrated with 2D animations and 3D structure-based movies

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English Journal

• C(16)-Ceramide-induced F-actin regulation stimulates mouse embryonic stem cell migration: Involvement of N-WASP/Cdc42/Arp2/3 complex and cofilin-1/α-actinin.

• Park SS, Kim MO, Yun SP, Ryu JM, Park JH, Seo BN, Jeon JH, Han HJ.SourceDepartment of Veterinary Physiology, Biotherapy Human Resources Center (BK 21), College of Veterinary Medicine, Chonnam National University, Gwangju 500-757, Republic of Korea.
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• Ceramide, a major structural element in the cellular membrane, is a key regulatory factor in various cellular behaviors that are dependent on ceramide-induced association of specific proteins. However, molecular mechanisms that regulate ceramide-induced embryonic stem cell (ESC) migration are still
• PMID 22989773

• Pathway of actin filament branch formation by Arp2/3 complex revealed by single-molecule imaging.

• Smith BA, Daugherty-Clarke K, Goode BL, Gelles J.SourceDepartment of Biochemistry, Brandeis University, Waltham, MA 02454-9110.
• Proceedings of the National Academy of Sciences of the United States of America.Proc Natl Acad Sci U S A.2013 Jan 22;110(4):1285-90. doi: 10.1073/pnas.1211164110. Epub 2013 Jan 4.
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• Doublecortin (dcx) family proteins regulate filamentous actin structure in developing neurons.

• Fu X, Brown KJ, Yap CC, Winckler B, Jaiswal JK, Liu JS.SourceCenter for Neuroscience Research and Center for Genetic Medicine Research, Children's National Medical Center, Washington, DC 20010, and Department of Neuroscience, University of Virginia Medical School, Charlottesville, Virginia 22908.
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Japanese Journal

• Localization of cortactin is associated with colorectal cancer development.

• Hirakawa Hiroshi,Shibata Kenichiro,Nakayama Toshiyuki
• International journal of oncology 35(6), 1271-1276, 2009-12
• … Cortactin is a ubiquitously expressed actin filament (F-actin)-binding protein that stabilizes F-actin networks and promotes actin polymerization by activating the actin-related protein 2/3 (Arp2/3) complex. …
• NAID 120002693653

• CCR7 Ligands Induce Endocytosis and the Formation of Actin-Filled Ruffles in Mature Dendritic Cells

• ,
• Journal of Clinical and Experimental Hematopathology 45(1), 25-35, 2005
• … In the present study, we examined effects of CCL19 and CCL21 on the organization of the actin cytoskeleton in murine DCs. … These chemokines considerably increased the intracellular level of fibrous actin and induced the formation of large petal-like ruffles of actin filaments in mature DCs. … In addition, a robust network of interconnected ruffles of actin filaments was assembled at the periphery of the cells after stimulation with the chemokines. …
• NAID 130000065674

• GRB2 links signaling to actin assembly by enhancing interaction of neural Wiskott-Aldrich syndrome protein (N-WASp) with actin-related protein (ARP2/3) complex

• CARLIER M. F.
• J Biol Chem 275, 21946-21952, 2000
• NAID 80011793878

Related Links

• ARPC2 - Actin-related protein 2/3 complex subunit 2 - Homo ...

Functions as actin-binding component of the Arp2/3 complex which is involved in regulation of actin polymerization and together with an activating nucleation-promoting factor (NPF) mediates the formation of branched actin networks.

• ATP hydrolysis on actin-related protein 2/3 complex causes ...

Fig. 1. ATP hydrolysis occurs on Arp2 after actin filament branching. (A) Photo-cross-linking of 8-azido-[γ-32 P]ATP to the Arp2/3 complex (2 μM) in the absence or presence of 20 μM WA. Samples were submitted to SDS/PAGE.

Related Pictures

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★リンクテーブル★

リンク元	「アクチン関連タンパク質 2/3複合体」
関連記事	「relate」「act」「complex」「complexed」「related protein」

「アクチン関連タンパク質 2/3複合体」

　　[★]

英

actin-related protein 2-3 complex

「relate」

　　[★]

• v.

• 関連する、関係する

関

associate、association、attach、bearing、concern、connect、connection、correlate、correlation、germane、implicate、implication、interplay、link、pertain、pertinent、referable、reference、relation、relationship、relative、relevance、relevant、respect

　 　 　 　

「act」

　　[★]

• v.

• 作用する、行動する、働く

• n.

• 行為、(法律)条例

関

action、behave、behavior、behavioral、behaviour、behavioural、conduct、effect、work

　 　 　 　 　 　 　 　 　 　 　 　 　 　

「complex」

　　[★]

• n.

• 複合体、錯体、複合物、合成物、(心理)コンプレックス

• adj.

• 複雑な、複合の

関

complexes、complicated、composite、conjugate、intricate

　 　 　 　 　

「complexed」

　　[★]

• adj.

• 複合型の、混合型の

関

hybrid、mixed lineage、mixed type、mixed-type

「related protein」

　　[★]

• 関連タンパク質、関連タンパク

関

associated protein