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Finalni Verze MASARYKOVA UNIVERZITA PŘÍRODOV ĚDECKÁ FAKULTA Ústav experimentální biologie Odd ělení genetiky a molekulární biologie Aktuální poznatky v molekulární biologii a epidemiologii viru ch řipky A Brno 2011 Bc. Silvie Tománková Pod ěkování Cht ěla bych pod ěkovat doc. RNDr. V. Růži čkové, CSc. za odborné vedení mé práce, poskytnutou literaturu, p řipomínky k textu a cenné rady b ěhem psaní této práce. 2 OBSAH Úvod ................................................................................................................................................4 1 Obecná charakteristika ............................................................................................................5 1.1 Virus ch řipky typu A .......................................................................................................5 2 Proteiny viru ch řipky typu A...................................................................................................6 2.1 Hemaglutinin (HA)..........................................................................................................6 2.2 Neuraminidáza (NA) .......................................................................................................6 2.3 Matrixový protein (M1)...................................................................................................7 2.4 Membránový protein (M2)..............................................................................................8 2.5 Nukleoprotein (NP) .........................................................................................................9 2.6 PB1-F2...........................................................................................................................10 2.7 RNA – dependentní RNA polymeráza..........................................................................10 2.7.1 Protein PA .............................................................................................................10 2.7.2 Protein PB1............................................................................................................11 2.7.3 Protein PB2............................................................................................................11 2.8 Nestrukturní fosfoprotein NS1 ......................................................................................11 2.9 Nestrukturní fosfoprotein NS2 ......................................................................................11 3 Reprodukce ch řipkového viru typu A ...................................................................................13 3.1.1 Aktivace polymerázového a transkrip čního komplexu .........................................14 3.1.2 Sbalování a pu čení virových částic .......................................................................14 4 Antigenní zm ěny ...................................................................................................................15 4.1 Antigenní drift (antigenní posun)..................................................................................16 4.2 Antigenní shift (antigenní zvrat)....................................................................................16 5 Ch řipkové onemocn ění..........................................................................................................18 5.1 Klinická manifestace .....................................................................................................18 5.2 Patogeneze.....................................................................................................................18 5.3 Lé čba .............................................................................................................................18 5.3.1 Blokátory iontového kanálu ch řipkového viru ......................................................19 5.3.2 Inhibitory virové neuraminidázy...........................................................................20 6 Pandemie ...............................................................................................................................20 6.1 Historický pohled na nejvýznamn ější pandemie ch řipky typu A..................................22 6.1.1 „Špan ělská ch řipka“ H1N1 z roku 1918................................................................22 6.1.2 „Asijská ch řipka“ H2N2 z roku 1957....................................................................22 6.1.3 Ch řipka „Hong Kong“ H3N2 z roku 1968............................................................23 6.1.4 „Pta čí ch řipka“ H5N1 (možný budoucí pandemický kmen).................................24 6.1.5 „Prase čí ch řipka“ H1N1 z roku 2009 ....................................................................24 7 Diagnostika ch řipkové infekce ..............................................................................................26 Záv ěr..............................................................................................................................................30 Seznam použitých zkratek.............................................................................................................31 Seznam použité literatury ..............................................................................................................33 3 Úvod Ch řipková infekce je často považována za banální onemocn ění, ale ve skute čnosti bývá příčinou mnoha úmrtí. Jsou známy t ři druhy p ůvodce této nemoci - Influenza A virus, Influenza B virus a Influenza C virus . Tyto obalené viry se segmentovaným RNA-genomem mají schopnost antigenní zm ěny, která jim pomáhá uniknout imunitnímu systému infikovaného hostitele. K přenosu této nákazy z člov ěka na člov ěka dochází kapénkovou infekcí s následným rychlým ší řením tohoto onemocn ění v sezónních epidemiích. Pouze ch řipka typu A m ůže vyvolat infekci v celosv ětovém m ěř ítku (tzv.pandemii), z d ůvodu vytvo ření nového sybtypu ch řipkového viru antigenním zvratem. Schopnost ch řipkového viru typu A zásadn ě zm ěnit svou genetickou informaci p ředstavuje celosv ětový zdravotnický problém. Cílem této práce je poskytnout základní informace týkající se aktuálních poznatk ů v molekulární biologii a epidemiologii viru ch řipky typu A se zvláštním z řetelem na ch řipkové onemocn ění lidí. 4 1 Obecná charakteristika Ch řipkové viry pat ří do čeledi Orthomyxoviridae. Jsou známy t ři druhy tohoto viru (Influenza A virus, Influenza B virus, Influenza C virus ), které se řadí do rod ů Influenzavirus A, Influenzavirus B a Influenzavirus C. Viry ch řipky byly objeveny ve dvacátém století. V roce 1933 byl izolován virus ch řipky A. W. Smithem, P. Laidlawem a C. H. Andersem. Virus ch řipky B objevil T. Jr. Francis v roce 1939 a virus ch řipky C byl objeven v roce 1950 R. M. Taylorem. Ch řipkové viry jsou obalené a uvnit ř kapsidu mají segmentovaný genom. To znamená, že negativní jedno řet ězcová nukleová kyselina RNA (-ssRNA) je rozd ělena na ur čitý po čet segment ů. Virus ch řipky typu A má stejn ě jako typ B osm segment ů, zatímco virus ch řipky C má segment ů sedm. 1.1 Virus ch řipky typu A Pr ůměrná velikost ch řipkového viru typu A se pohybuje okolo 120 nm (proto ho lze pozorovat pouze v elektronovém mikroskopu) a velikost genomu je p řibližn ě 15 kb. Tento virus má dva povrchové glykoproteinové antigeny – hemaglutinin (HA) a neuraminidázu (NA). Geny lokalizované na osmi segmentech genomu kódují 11 genových produkt ů, mezi které pat ří (viz. Obr. 1): 1) proteiny polymerázového komplexu: PB1, PB1-F2, PB2 a PA (funkce RNA transkriptázy a replikázy) 2) proteiny: HA (hemaglutinin), NP (nukleoprotein), NA (neuraminidáza), M1 a M2 (matrixové proteiny), NS1 a NS2 (nestrukturní proteiny) Nomenklatura ch řipkových vir ů typu A je rozd ělena do jednotlivých subtyp ů podle jejich povrchových antigen ů (glykoprotein ů) na povrchu virionu hemaglutininu (H1 - H16) a neuraminidázy (N1 - N9). 5 Obrázek 1: Struktura virionu ch řipky typu A (http://www.nature.com/scitable/topicpage/genetics-of-the-influenza- virus-716) 2 Proteiny viru ch řipky typu A 2.1 Hemaglutinin (HA) Je membránový protein a hlavní povrchový antigen viru ch řipky typu A. Tento protein zodpovídá za vazbu k receptor ům hostitelské bu ňky (jako receptor slouží kyselina sialová) a za fúzi mezi virionem a hostitelskou bu ňkou (uvoln ění nukleokapsidu do cytoplazmy bu ňky). Aktivní je v podob ě trimeru a je kódován čtvrtým RNA segmentem. Podléhá t řem r ůzným posttransla čním proces ům (proteolytickému št ěpení, glykosylaci a acylaci mastných kyselin). Nov ě syntetizovaný HA je št ěpen, a to tak, že je odstran ěno 14 - 18 AMK, které slouží jako signální sekvence pro jeho transport k bun ěč né membrán ě (Webster et al. , 1992; Isin et al. , 2002). Proteolytickým štěpením HA vznikají dv ě podjednotky (HA1 a HA2), které jsou propojeny disulfidickými vazbami. Toto št ěpení je zp ůsobeno proteázami, které jsou produkovány hostitelskou bu ňkou a jsou podobné trypsinu. Tento d ěj je podmínkou infek čnosti, jelikož fúze viru a bu ňky je zprost ředkována volným amino-koncem HA2. Nešt ěpený HA (HA0) se tedy skládá z HA1 (obvykle 324 AMK + sacharid) a HA2 (obvykle 222 AMK + sacharid + 3 zbytky kyseliny palmitové). B ěhem maturace tvo ří HA homotrimery. V podstat ě každá molekula HA se skládá z globulární hlavy na stopce . Hlava je tvo řena výhradn ě podjednotkou HA1 a je schopná vázat receptor, zatímco stopka je složena z HA2 a částe čně i z HA1. 6 Karboxy-terminální oblast HA2 obsahuje hydrofobní transmembránové sekvence a terminální − cytoplazmatickou sekvenci, kde je p řipojen palmitát /CH 3(CH 2)14
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