WordNet

the act of attacking; "attacks on women increased last year"; "they made an attempt on his life" (同)attempt
(military) an offensive against an enemy (using weapons); "the attack began at dawn" (同)onslaught, onset, onrush
a decisive manner of beginning a musical tone or phrase (同)tone-beginning
an offensive move in a sport or game; "they won the game with a 10-hit attack in the 9th inning"
strong criticism; "he published an unexpected attack on my work"
a sudden occurrence of an uncontrollable condition; "an attack of diarrhea"
begin to injure; "The cancer cells are attacking his liver"; "Rust is attacking the metal"
set to work upon; turn ones energies vigorously to a task; "I attacked the problem as soon as I got out of bed"
the onset of a corrosive or destructive process (as by a chemical agent); "the film was sensitive to attack by acids"; "open to attack by the elements"
attack in speech or writing; "The editors of the left-leaning paper attacked the new House Speaker" (同)round, assail, lash out, snipe, assault
take the initiative and go on the offensive; "The Serbs attacked the village at night"; "The visiting team started to attack" (同)aggress
launch an attack or assault on; begin hostilities or start warfare with; "Hitler attacked Poland on September 1, 1939 and started World War II"; "Serbian forces assailed Bosnian towns all week" (同)assail
make complete or perfect; supply what is wanting or form the complement to; "I need some pepper to complement the sweet touch in the soup"
a complete number or quantity; "a full complement"
one of a series of enzymes in the blood serum that are part of the immune response
something added to complete or embellish or make perfect; "a fine wine is a perfect complement to the dinner"; "wild rice was served as an accompaniment to the main dish" (同)accompaniment
number needed to make up a whole force; "a full complement of workers" (同)full complement
a word or phrase used to complete a grammatical construction
either of two parts that mutually complete each other
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"
a compound described in terms of the central atom to which other atoms are bound or coordinated (同)coordination_compound
a conceptual whole made up of complicated and related parts; "the complex of shopping malls, houses, and roads created a new town" (同)composite
(psychoanalysis) a combination of emotions and impulses that have been rejected from awareness but still influence a persons behavior
a thin pliable sheet of material
a pliable sheet of tissue that covers or lines or connects the organs or cells of animals or plants (同)tissue layer

PrepTutorEJDIC

…'を'『襲う』,『攻撃する』;(ゲームで)…'を'攻める / (演説・文書などで)…'を'『非難する』,攻撃する / 〈病気・不運などが〉…'を'襲う,冒す / 〈仕事など〉‘に'勢いよく取りかかる,着手する / 『攻撃をする』 / 〈U〉〈C〉(…に対する)『攻撃』;(ゲームの)攻め;非難《+『on』(『upon, against』)+『名』》 / 〈C〉発病,発作 / 〈U〉(仕事への)着手,開始
(あるものを完全にするため)(…を)補う物(事)《+『of』+『名』》 / 補語(文法で文の成分の一つ) / (必要な)全数,全量;(船の)定員 / …を補足する,補う
『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)
(生物の)膜,皮膜

Wikipedia preview

出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2017/05/11 15:18:02」(JST)

wiki en

[Wiki en表示]

Membrane Attack Complex (Terminal Complement Complex C5b-9)

A membrane attack complex attached to a pathogenic cell.

The membrane attack complex (MAC) or terminal complement complex (TCC) is a structure typically formed on the surface of pathogenic bacterial cells as a result of the activation of the host's alternative pathway, classical pathway, or lectin pathway of the complement system, and it is one of the effector proteins of the immune system. The membrane-attack complex (MAC) forms transmembrane channels. These channels disrupt the cell membrane of target cells, leading to cell lysis and death.^[1]^[2]

A number of proteins participate in the assembly of the MAC. Freshly activated C5b binds to C6 to form a C5b-6 complex, then to C7 forming the C5b-6-7 complex. The C5b-6-7 complex binds to C8, which is composed of three chains (alpha, beta, and gamma), thus forming the C5b-6-7-8 complex. C5b-6-7-8 subsequently binds to C9^[3]^[4]^[5] and acts as a catalyst in the polymerization of C9.

Active MAC has a subunit composition of C5b-C6-C7-C8-C9{n}.

Structure and function

It is composed of a complex of four complement proteins (C5b, C6, C7, and C8) that bind to the outer surface of the plasma membrane, and many copies of a fifth protein (C9) that hook up to one another, forming a ring in the membrane. C6-C9 all contain a common MACPF domain.^[6] This region is homologous to cholesterol-dependent cytolysins from Gram-positive bacteria.^[7]

The ring structure formed by C9 is a pore in the membrane that allows free diffusion of molecules in and out of the cell. If enough pores form, the cell is no longer able to survive.

If the pre-MAC complexes of C5b-7, C5b-8 or C5b-9 do not insert into a membrane, they can form inactive complexes with Protein S (sC5b-7, sC5b-8 and sC5b-9). These fluid phase complexes do not bind to cell membranes and are ultimately scavenged by clusterin and vitronectin, two regulators of complement.^[8]

Initiation: C5-C7

Membrane attack complex.

The membrane attack complex is initiated when the complement protein C5 convertase cleaves C5 into C5a and C5b.

Another complement protein, C6, binds to C5b.

The C5bC6 complex is bound by C7.

This junction alters the configuration of the protein molecules exposing a hydrophobic site on C7 that allows the C7 to insert into the phospholipid bilayer of the pathogen.

Polymerization: C8-C9

Similar hydrophobic sites on C8 and C9 molecules are exposed when they bind to the complex, so they can also insert into the bilayer.

C8 is a complex made of the two proteins C8-beta and C8 alpha-gamma.

C8 alpha-gamma has the hydrophobic area that inserts into the bilayer. C8 alpha-gamma induces the polymerization of 10-16 molecules of C9 into a pore-forming structure known as the membrane attack complex.

It has a hydrophobic external face allowing it to associate with the lipid bilayer.
It has a hydrophilic internal face to allow the passage of water.

Multiple molecules of C9 can join spontaneously in concentrated solution to form polymers of C9. These polymers can also form a tube-like structure.

Inhibition

CD59 acts to inhibit the complex. This exists on body cells to protect them from MAC. A rare condition, paroxysmal nocturnal haemoglobinuria, results in red cells that lack CD59. These red cells can, therefore, be lysed by MAC.

Pathology

Deficiencies of C5 to C9 components does not lead to generic infections, but only to increased susceptibility to Neisseria spp.,^[9] since these bacteria have a thin cell wall and glycocalix.^[10]

References

^ Peitsch MC, Tschopp J (1991). "Assembly of macromolecular pores by immune defense systems". Curr. Opin. Cell Biol. 3 (4): 710–716. doi:10.1016/0955-0674(91)90045-Z. PMID 1722985.
^ Reid K. B. M., The complement system, in: B. D. Hames and D. M. Glover (eds.), Molecular Immunology, Oxford: IRL Press, 1988, pp. 189-241.
^ Stanley KK, Marazziti D, Eggertsen G, Fey GH (1988). "Relationships between the gene and protein structure in human complement component C9". Biochemistry. 27 (17): 6529–6534. doi:10.1021/bi00417a050. PMID 3219351.
^ Stanley KK, Luzio JP, Tschopp J, Kocher HP, Jackson P (1985). "The sequence and topology of human complement component C9". EMBO J. 4 (2): 375–382. PMC 554196. PMID 4018030.
^ Fey GH, Hugli TE, Podack ER, Gehring MR, Kan CC, DiScipio RG (1984). "Nucleotide sequence of cDNA and derived amino acid sequence of human complement component C9". Proc. Natl. Acad. Sci. U.S.A. 81 (23): 7298–7302. doi:10.1073/pnas.81.23.7298. PMC 392133. PMID 6095282.
^ Tschopp J, Masson D, Stanley KK (1986). "Structural/functional similarity between proteins involved in complement- and cytotoxic T-lymphocyte-mediated cytolysis". Nature. 322 (6082): 831–4. doi:10.1038/322831a0. PMID 2427956.
^ Carlos J. Rosado; Ashley M. Buckle; Ruby H. P. Law; Rebecca E. Butcher; Wan-Ting Kan; Catherina H. Bird; Kheng Ung; Kylie A. Browne; Katherine Baran; Tanya A. Bashtannyk-Puhalovich; Noel G. Faux; Wilson Wong; Corrine J. Porter; Robert N. Pike; Andrew M. Ellisdon; Mary C. Pearce; Stephen P. Bottomley; Jonas Emsley; A. Ian Smith; Jamie Rossjohn; Elizabeth L. Hartland; Ilia Voskoboinik; Joseph A. Trapani; Phillip I. Bird; Michelle A. Dunstone & James C. Whisstock (2007). "A Common Fold Mediates Vertebrate Defense and Bacterial Attack". Science. 317 (5844): 1548–51. doi:10.1126/science.1144706. PMID 17717151.
^ Hadders, MA (2012). "Assembly and regulation of the membrane attack complex based on structures of C5b6 and sC5b9". Cell Rep. 1. doi:10.1016/j.celrep.2012.02.003.
^ Ronald Hoffman, Leslie E. Silberstein, Helen Heslop, Jeffrey Weitz, Hematology: Basic Principles and Practice, 6th ed., Elsevier, 2013, page 231.
^ Abul K. Abbas, Andrew H. H. Lichtman, Shiv Pillai, Basic Immunology: Functions and Disorders of the Immune System, 5th ed., Elsevier, 2015, page 179.

External links

Complement Membrane Attack Complex at the US National Library of Medicine Medical Subject Headings (MeSH)
http://www.lib.mcg.edu/edu/esimmuno/ch2/lysis.htm

UpToDate Contents

全文を閲覧するには購読必要です。 To read the full text you will need to subscribe.

1. 補体系の概要および臨床的評価 overview and clinical assessment of the complement system
2. 補体系の制御因子および受容体 regulators and receptors of the complement system
3. 補体経路 complement pathways
4. 補体系の遺伝性疾患 inherited disorders of the complement system
5. 糸球体の免疫障害の機序 mechanisms of immune injury of the glomerulus

English Journal

The membrane attack complex as an inflammatory trigger.

Morgan BP1.
Immunobiology.Immunobiology.2016 Jun;221(6):747-51. doi: 10.1016/j.imbio.2015.04.006. Epub 2015 Apr 30.
The final common pathway of all routes of complement activation involves the non-enzymatic assembly of a complex comprising newly formed C5b with the plasma proteins C6, C7, C8 and C9. When assembly occurs on a target cell membrane the forming complex inserts into and through the bilayer to create a
PMID 25956457

Soluble CD59 is a Novel Biomarker for the Prediction of Obstructive Chronic Lung Allograft Dysfunction After Lung Transplantation.

Budding K1, van de Graaf EA2, Kardol-Hoefnagel T1, Kwakkel-van Erp JM2, Luijk BD2, Oudijk EJ3, van Kessel DA3, Grutters JC3, Hack CE1,4, Otten HG1.
Scientific reports.Sci Rep.2016 May 24;6:26274. doi: 10.1038/srep26274.
CD59 is a complement regulatory protein that inhibits membrane attack complex formation. A soluble form of CD59 (sCD59) is present in various body fluids and is associated with cellular damage after acute myocardial infarction. Lung transplantation (LTx) is the final treatment for end-stage lung dis
PMID 27215188

Complement system activation contributes to the ependymal damage induced by microbial neuraminidase.

Granados-Durán P1, López-Ávalos MD1, Hughes TR2, Johnson K3, Morgan BP2, Tamburini PP3, Fernández-Llebrez P1, Grondona JM4.
Journal of neuroinflammation.J Neuroinflammation.2016 May 21;13(1):115. doi: 10.1186/s12974-016-0576-9.
BACKGROUND: In the rat brain, a single intracerebroventricular injection of neuraminidase from Clostridium perfringens induces ependymal detachment and death. This injury occurs before the infiltration of inflammatory blood cells; some reports implicate the complement system as a cause of these inju
PMID 27209022

Japanese Journal

補体の沈着を認めなかったPLA2R 陽性特発性膜性腎症の1 例

日本小児腎臓病学会雑誌 29(1), 33-38, 2016
NAID 130005145034

補体の沈着を認めなかったPLA<sub>2</sub>R 陽性特発性膜性腎症の1 例

日本小児腎臓病学会雑誌 advpub(0), 2016
NAID 130005125666

Current treatment of atypical hemolytic uremic syndrome

Intractable & Rare Diseases Research 3(2), 34-45, 2014
NAID 130005008084

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リンク元	「膜侵襲複合体」
関連記事	「attack」「complex」「complement」「complexed」「complexes」

「膜侵襲複合体」

　　[★]

英: membrane attack complex, MAC
同: 膜傷害複合体
関: complement membrane attack complex

[show details]

膜攻撃複合体 : 76 件
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溶菌作用を有する補体カスケード産物C5b6789複合体

「attack」

　　[★]

n.

発作、攻撃

v.

攻撃する、襲う、非難する、襲撃する

関: aggressive、assault、blame、bout、challenge、crises、crisis、criticise、criticism、criticize、insult、offense、offensive、paroxysm、seizure、strike、stroke、thrust

「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

「complement」

　　[★]

n.

補って完全にする物、補完物。(免疫)補体
(必要な)全数、善良

vt.

補って完全にする。補完する。～の補足となる。

「complexed」

　　[★]

adj.

複合型の、混合型の

関: hybrid、mixed lineage、mixed type、mixed-type

「complexes」

　　[★]

n.

複合体

関: complex、composite

[PUB00001056-1] Peitsch MC, Tschopp J (1991). "Assembly of macromolecular pores by immune defense systems". Curr. Opin. Cell Biol. 3 (4): 710–716. doi:10.1016/0955-0674(91)90045-Z. PMID 1722985.

[2] Reid K. B. M., The complement system, in: B. D. Hames and D. M. Glover (eds.), Molecular Immunology, Oxford: IRL Press, 1988, pp. 189-241.

[PUB00000295-3] Stanley KK, Marazziti D, Eggertsen G, Fey GH (1988). "Relationships between the gene and protein structure in human complement component C9". Biochemistry. 27 (17): 6529–6534. doi:10.1021/bi00417a050. PMID 3219351.

[PUB00001141-4] Stanley KK, Luzio JP, Tschopp J, Kocher HP, Jackson P (1985). "The sequence and topology of human complement component C9". EMBO J. 4 (2): 375–382. PMC 554196. PMID 4018030.

[PUB00004608-5] Fey GH, Hugli TE, Podack ER, Gehring MR, Kan CC, DiScipio RG (1984). "Nucleotide sequence of cDNA and derived amino acid sequence of human complement component C9". Proc. Natl. Acad. Sci. U.S.A. 81 (23): 7298–7302. doi:10.1073/pnas.81.23.7298. PMC 392133. PMID 6095282.

[6] Tschopp J, Masson D, Stanley KK (1986). "Structural/functional similarity between proteins involved in complement- and cytotoxic T-lymphocyte-mediated cytolysis". Nature. 322 (6082): 831–4. doi:10.1038/322831a0. PMID 2427956.

[macpfstructure-7] Carlos J. Rosado; Ashley M. Buckle; Ruby H. P. Law; Rebecca E. Butcher; Wan-Ting Kan; Catherina H. Bird; Kheng Ung; Kylie A. Browne; Katherine Baran; Tanya A. Bashtannyk-Puhalovich; Noel G. Faux; Wilson Wong; Corrine J. Porter; Robert N. Pike; Andrew M. Ellisdon; Mary C. Pearce; Stephen P. Bottomley; Jonas Emsley; A. Ian Smith; Jamie Rossjohn; Elizabeth L. Hartland; Ilia Voskoboinik; Joseph A. Trapani; Phillip I. Bird; Michelle A. Dunstone & James C. Whisstock (2007). "A Common Fold Mediates Vertebrate Defense and Bacterial Attack". Science. 317 (5844): 1548–51. doi:10.1126/science.1144706. PMID 17717151.

[8] Hadders, MA (2012). "Assembly and regulation of the membrane attack complex based on structures of C5b6 and sC5b9". Cell Rep. 1. doi:10.1016/j.celrep.2012.02.003.

[9] Ronald Hoffman, Leslie E. Silberstein, Helen Heslop, Jeffrey Weitz, Hematology: Basic Principles and Practice, 6th ed., Elsevier, 2013, page 231.

[10] Abul K. Abbas, Andrew H. H. Lichtman, Shiv Pillai, Basic Immunology: Functions and Disorders of the Immune System, 5th ed., Elsevier, 2015, page 179.

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案内

complement membrane attack complex