ࡱ>  '`BbjbjLULU .?.?A&x`x`x`8`,`q$2lbb"bbbKgKgKg"""""""$%h (p"Yhf^Kghh"bbI+$h8bb"h"B`b`b M^x`} $fA$0q$N{({({( KgLg6g,gKgKgKg""^KgKgKgq$hhhhdXx`x`  1. UVOD Rak je klonalna bolest koja nastaje greakom u regulaciji stani nih mehanizama proliferaciji, diferencijaciji, i stani nom pre~ivljavanju. To je bolest koja nastaje iz jedne stanice koja do~ivi svoju prvu mutaciju uslijed ega se sposobna br~e umna~ati, a time i mutirati. Pojavom sljedee mutacije jedna od novonastalih stanica e joa br~e proliferirati i u njoj e se joa br~e odvijati mutacijske promjene. Ciklus se tako nastavlja, a mutirane stanice sve viae dobivaju obilje~ja prave kancerogene stanice, stanice sposobne za brzu proliteraciju, gotovo neovisno o faktorima rasta, sposobne za napad na zdravo tkivo i na kraju sposobne za irenje iz primarnog sredita razvoja raka i metastaziranje u druge dijelove tijela dostupne krvi, naruaavajui normalnu funkciju tkiva i organa.1 Nekontrolirano umna~anje kancerogenih stanica posljedica je napravilnog odgovora na signale iz svoje okoline zbog nakupljenih abnormalnosti/mutacija u viae regulacijskih sustava stanice. Kancerogene stanice ne odgovaraju na signale za inhibiciju proliferacije i imaju smanjenu potrebu za faktorima rasta. Zahvaljujui svoji promjenjenim signalnim putevima, te stanice odgovaraju punim intezitetom na signale iz svoje okoline koji su ina e preslabi da bi aktivirali normalne stanice, ili pretjerano odgovaraju na signale normalne ja ine. Kancerogene stanice esto nisu dovoljno diferencirane, ato je u direktnoj vezi s poveanom proliferacijom tih stanica.Kancerogene stanice esto zaobilaze mehanizam programirane stani ne smrti (apoptoze) produljujui svoj ~ivotni vijek, nakupljajui joa viae mutacija i time razvijajui joa maligniji fenotip, ato znatno doprinosi razvoju raka. William Dameshek je 1951. godine primjetio da ve otprije poznate bolesti, tada znane pod nazivima kroni na granulocitna leukemija, policitemija, idiopatska mijeloidna metaplazija, trombocitemija, megakariocitna leukemija i eritroleukemija, imaju zajedni ke karakteristike unato  naizgled velikoj varijabilnosti u simptomima, krvnoj slici i na inu napredovanja bolesti.Pretpostavio je da su sve one zapravo istog postanka-preintezivne proliferacije koatane sr~i i da se svrstaju u jednu zajedni ku kategoriju koju je nazvao mijeloproliferativni poremeaji (MPD). Kasnije su ovi poremeaji podjeljeni na nasljedne (u kojima uvijek dolazi do istih mutacija u roditelja i u potomaka koji direktno uzrokuju bolesti) i obiteljski skolone ili prave mijeloproliferativne poremeaje (u kojima se nasljeuje samo sklonost razvijanju razli itih mutacija koje zajedno uzrokuju razvoj odreenog poremeaja).2 Otkriveno je da se u potonjima esto pronalazi somatska mutacija gena za citoplazmatsku kinazu JAK2, V617F, te da ona vjerojatno igra neku od zna ajnijih uloga u razvitku MPD.3 1.1 Mijeloproliferativni poremeaji Mijeloproliferativne bolesti (MPB) su klonalni poremeaji mati ne krvotvorne stranice koje se o ituju poja anjim stvaranjem i nakupljanjem velikog broja krvnih stranica: eritrocita, granulocita i trombocita, u koatanoj sr~i, perifernoj krvi i nekim tkivima. esto se zbog poveanog broja stanica u krvi mogu razviti mikrovaskularne smetnje, ali i ozbiljnije tromboze. Nedavna studija je pokazala da viae od treine pacijenata koji boluju od tromboze (Budd-Chiari sindrom, tromboza portalne ili mezenteri ne vene) ima neki mijeloproliferativni poremeaj.4 Zbog klini ki vrlo raznolikih svojstava (razina hemoglobina, broj krvnih stanica, splenomegalija) mijeloproliferativni poremeaji su ispo etka bili podjeljeni na brojne skupine koje su meusobno essto znale imati sli ne stadije bolesti. Svetska zdravstvena organizacija (WHO) je stoga predlo~ila kriterije za precizniju dijagnostiku zasnovanu na patoloakim obilje~jima bolesti (analiza koatane sr~i) (WHO 2001). Iako je klasifikacija donijela red u dijagnostiku mijeloproliferativnih poremeaja, tek je zajedni kim naporom sljedea skupina znanstvenika zaista napravila pomak u klasifikaciji integrirajui klini ke i patoloake, ali i molekularne markere pod zajedni kim nazivom Europski klini ki i patoloaki (ECP) kriterij.2 Prema klini kim svojstvima, molekularnom mehanizmu nastanka bolesti i na inu lije enja bilo je o igledno da se kroni na mijeloi na leukemija (CML) prili no razlikuje od ostatka mijeloproliferativnih poremeaja (MPD). CML karakterizira nakupljanje zrelih i nezrelih oblika granulocita u koatanoj sr~i i perifernoj krvi. Za CML je karakteristi na specifi na citogenetska promjena Philidelphia kromosom. To je recipro na translokacija razli ite koli ine genetskog materijala s dugog kraka kromosoma 22 (22q-) na drugi kromosom, u viae od 90 % slu ajeva na dugi krak kromosoma 9. Takva se promjena ozna ava t(9q+:22q-). Za razliku od CML mehanizam nastanka mijeloproliferativnih poremeaja je nepoznat. MPD je vrlo heterogena i varijabilna, i zbog toga relativno teako izlje iva skupina bolesti. Prema ECP kriterijima mijeloproliferacije se dijele u tri glavna poremeaja: policitemija rubra vera (PRV) esencijalna trombocitemija (ET) kroni na idiopatska mijelofibroza (CIMF) 1.1.1. Policitemija rubra vera (PRV) PRV je neoplasti na klonalna mijeloproliferativna bolest nastala transformacijom pluripotentne mati ne stanice. Klini ki se o ituje hiperplazijom hematopoeze u koatanoj sr~i zbog nekontrolirane proliferacije eritrocitopoeze, granulocitopoeze i trombocitopoeze. U perifernoj krvi dominira eritrocitoza, a nalazi se i umjerena leukocitoza i trombocitoza. Kroni nog je i progresivnog tijeka te u kasnijoj fazi esto ulazi u tzv. fazu iscrpljenje koja odgovara slici idiopatske mijelofibroze. U estalost bolesti je 0,5 - 2 na 100 000 stanovnika. Naj eae se dijagnosticira u aestom i sedmom desetljeu ~ivota, neato eae u muakaraca nego ~ena. Etiologija bolesti nije poznata, ali je sigurno da bolest nije posljedica prekomjerna lu enja ili osjetljivosti stanica na eritropoetin. Smetnje koje PRV uzrokuje su naj eae posljedica poveane viskoznosti krvi i sklonosti tromboembolijama. Venske tromboze se naj eae javljaju na donjim udovima, a arterijske tromboembolije se o ituju cerebrovaskularnom ili koronarnom simptomatologijom. Pri pregledu bolesnik je obi no pletori an, zagasito crvene boje ko~e i vidljivih sluznica. Sedimentacija eritrocita je sni~ena, a koncentracija hemoglobina i hematokrit su poviaeni. U pravilu se nalazi arterijska hipertenzija, a nerijetko i splenomegalija. Cilj lije enja je sprije iti pojavu tromboembolijskih komplikacija i smanjiti simptome bolesti. To se mo~e postii smanjenjem broja eritrocita (citoredukcija) i antiagregacijskim lije enjem. Citoredukcija se provodi venepunkcijom i primjenom citostatika (hidroksiureja), a antiagregacijsko lije enje se provodi malim dozama acetilsalicilne kiseline. Pravilno lije eni PRV je kroni na bolest koja samo donekle skrauje ~ivotni vijek bolesnika. 1.1.2. Esencijalna trombocitemija (ET) ET je klonalni poremeaj multipotentne krvotvorne mati ne stanice, a malignom klonu pripadaju sve krvne loze. Nejasno je zaato proliferira samo trombocitopoeza. Abnormalno je proairen odjeljak megakariocitnih mati nih stanica u koatanoj sr~i, ato nije slu aj kod stanja sa sekundarnom trombocitozom. ET rijetko izaziva ope simptome tipi ne za druge mijeloproliferativne bolesti, na primjer umor, znojenje, gubitak te~ine i sl. esto se otkriva slu ajem pregledom krvne slike. U krvnoj slici se nalazi trombocitoza, mo~e postojati leukocitoza na ra un granulocitoze a crvena krvna slika je normalna. Naj eae komplikacije bolesti su arterijske tromboembolije koje se o ituju cerebrovaskularnim i koronarnim simptomima. U bolesnika s izrazitim poviaenjem broja trombocita (preko 1000 x 109.) dolazi do poja ane sklonosti krvarenju. Bolesnici obi no krvare u ko~u ili vidljive sluznice ato, iako katkad izgleda dramati no, u velikoj veini slu ajeva nije opasno. Iako nema sigurnih dokaza da sni~enje broja trombocita utje e na prognozu asimptomatskih bolesnika sa ET, logi an je cilj terapije sniziti broj trobmocita. Za citoredukciju se naj eae rabe hidroksiureja i interferon-alfa, a acetilsalicilna kiselina se koristi u bolesnika koji imaju opetovane tromboti ke komplikacije. U usporedbi s opom populacijom o ekivani ~ivotni vijek bolesnika s ET-om nije zna ajnije skraen. 1.1.3. Kroni na idiopatska mijelofibroza (CIMF) CIMF je klonalna mijeloproliferativna bolest koju karakterizira ekstramedularna hematopoeza i izrazita fibroza koatane sr~i. Mijeloproliferacija zahvaa prvenstveno granulocitnu i megakariocitnu lozu s posljedi nom leukocitozom i trombocotiozom. Iako je najprominentniji nalaz fibroza koatane sr~i, fibroblasti nisu dio malignog klona i njihovo je bujanje i nastanak fibroze sekundarna pojava zbog poveanog izlu ivanja faktora rasta koji stimuliraju fibroblasti. Uzrok bolesti je nepoznat, a javlja se obi no u dobi iznad 50. godine s jednakom u estaloau oba spola. Nespecifi ni simptomi i znakovi kao ato su brzo umaranje, slabost, dispnea i gubitak te~ine prisutni su u veine bolesnika. Splenomegalija je est simptom, a bol u podru ju slezene ili lijevom ramenu govori u prilog infarktu slezene. Karakteristi an nalaz su divovski trombociti i njihova nenormalna granulacija te nalaz golih jezgara megakariocita. Biopsija koatane sr~i je najva~niji dijagnosti ki postupak, a aspiracija koatane sr~i zbog fibroze naj eae ne uspijeva. Nema terapijskog pristupa koji bitno produ~ava ~ivot bolsenika s CIMF. U mladih bolesnika koji imaju histokompatibilnog srodnika valja razmotriti lije enje transplatacijom koatane sr~i. Medijan pre~ivljenja iznosi 4 do 5 godina od dijagnoze a tijek je kroni an i progresivan; u oko 10% slu ajeva javlja se akutna mijeloi na leukemija. Naj eai uzroci smrti su infekcije, krvarenja, transformacija u akutnu leukemiju, zatajivanje srca ili bubrega te portalna hipertenzija.5 1.2. Mijeloproliferativni poremeaji i JAK2 Glavna odlika mijeloproliferativnih poremeaja jest da su hematopoetske progenitorne stanice preosjetljive na mnoge faktore rasta i druge citokine zbog kojih one ubrzano rastu i proliferiraju. Ipak, broj receptora za ove signalne molekule na povrini stanica je nepromijenjen to ukazuje da se glavna greka nalazi unutar ovih stanica, u signalnom putu koji je u estoj i airokoj uporabi. Fosforilacija i defosforilacija su osnovni regulativni mehanizmi rasta i proliferacije stanica. Mutacije protein tirozin kinaza i fosfataza mogu se pronai u mnogim ljudskim tumorima, a logi na je pretpostavka da su one uzrokom i mijeloproliferativnih poremeaja.6 Pet je istra~iva kih grupa gotovo istodobno otkrilo mutaciju JAK2 (V617F) u pacijenata oboljelih od mijeloproliferativnih poremeaja. Baxter i sur. su ispitivali mutacije u tirozin kinazama i detektirali u estalu to kastu mutaciju u visoko konzerviranom slijedu egzona 12 gena jak2.3 Ova G-T transverzija na nukleotidu 1849 je uzrokovala zna ajnu promjenu na JAK2 proteinu zamjenivai valin s mjesta 617 za fenilalanin u JH2 domeni. Kralovics i sur. su citogeneti kim metodama ispitivali kromosomske aberacije, i manje mutacije kao, na primjer, gubitak heterozigotnosti 8LOH po cijelom genomu granulocita PRV pacijenata. Kromosomske aberacije nisu pronaene, ali je pojava LOH negdje na kraem (p) kraku kromosoma 9 bila dosta esta. LOH nastje mitotskom rekombinacijom telomerne regije homolognog para kromosom 9. Ispitujui samo 23 gena koja su aktivna u multipotentnim hematopoetskim stanicama CD34+ na vrhu 9p kraka, detektirana je to kasta mutacija na genu jak2.7 James i sur. su provjeravali osjetljivost hematopoetskih mati nih stanica PRV pacijenata u kulturi na inhibitore kinaza i otkrili dva inhibitora koja sprije avaju spontani rast eritroidnih mati nih stanica (EEC) karakteristi an za PRV. Oba inhibitora su djelovala na JAK2. Sekvecioniranjem gena jak2 pronaali su JAK2 (V617F) mutaciju.8 Levine i sur. i Zhao i sur. su pretpostavili da su svi MPD uzrokovani mutacijama u tirozin kinazama, jednako kao ato je CML uzrokovana mutacijom u tirozin kinazi. Stoga su sekvencirali sve tirozin kinaze u genomskoj DNA pacijenata s PRV i pronaali istu mutaciju JAK2 (V617F).9 Ova mutacija je preko noi postala jedan od glavnih kriterija dijagnostike mijeloproliferativnih poremeaja, ali i primarna meta u daljnim istra~ivanjima MPD i razvoju lije enja.3 Vainchenker je iznio pretpostavku da je uzrok PRV neminovno povezan sa ovom mutacijom,ali mehanizam nastanka bolesti je i dalje ostao nejasan. 8 1.2.1. Struktura i funkcija Janus kinaze 2 Janus kinaze su otkrivene 1990. godine kao tada jedinstvena skupina citoplazmatskih protein tirozin kinaza ija je uloga prijenos signala sa povraine stanice prema jezgri. Nazvane su prema izrazu samo joa jedna kinaza (engl. Just Another Kinase JAK), ali ubrzo nakon to je definirana njihova iroka uloga u prijenosu signala citoplazmom, promijenjeno im je ime u Janus kinase (prema starorimskom bogu s dva lica, Ianusu). JAK2 ima vie razina regulacije aktivnosti. Prva je fosforilacija aktivacijske petlje kinazne domene koja aktivira kinazu. Druga razina je esteri ka inhibicija pseudokinazne domene nad kinaznom. Trea razina regulacije se zasniva na fosforilaciji FERM domene koja ina e ve~e JAK2 za citoplazmatski dio receptora blizu membrane, omoguava dimerizaciju kinaze i tako je aktivira. Fosforilacijom FERM domene, JAK2 disocira s receptora i razgrauje se.10 1.2.2 JAK2 (V617F) mutacija Mutacije Janus kinaza u leukemijama i hemapoetskim sindromima nisu novost. Najpoznatija mutacija na JAK2 su kromosomske translokacije iz kojih nastaju fuzije Janus kinaze 2 sa drugim proteinima i kinazama. Mutacija JAK2 (V617F) se nalazi u veini mijeloidnih stanica veine pacijenata. Ona stoga mora davati neku selektivnu prednost mutiranoj stanici naspram zdravih stanica i tako doprinijeti patogenezi mijeloproliferativnih poremeaja. Eritroidne mati ne stanice, koje nose ovu mutaciju rastu u kulturi bez eritroopoetina. To meutim nije slu aj i kod normalnih stanica, pa se mo~e zaklju iti da mutacija JAK2 (V617F) uzrokuje neovisnost stani nog rasta i proliferacije od faktora rasta. Jedno od objaanjenja za ovakvo ponaaanje jest da mutacija JAK2 (V617F) odr~ava Janus kinazu 2 stalno aktivnom, bez obzira je li eritropoetin prisutan ili ne. Budui da je izrazito rijetka u T- limfocitima, za pretpostaviti je da je mutacija V617F na JAK2 ste ena. Kako je ta mutacija prisutna i u granulocitno-makrofagnoj i u eritroidnoj hematopoetskoj lozi, morala je nastati ve u multipotentnoj hematopoetskoj mati noj stanici, pretku obiju loza.3 Ipak, ne smije se zanemariti mogunost da je dio mutirane populacije stanica mogao nastati i u diferenciranijim hematopoetskim stanicama.10 Dovoljno je da samo jedan alel jak 2 bude mutiran i da se JAK2 (V617F) konstitiutivno aktivira i potakne proliferaciju megakariocita. Kotransfekcijskim eksperimentima dokazano je da su JAK2 i JAK2 (V617F) konkurenti za iste signalne puteve, pa bi gubitak i drugog gena jak2 divljeg tipa dao znatnu selektivnu prednost stanici.8 Ali fosforilacijski eksperimenti pokazuju da je samo jedan mutirani alel jak2 dovoljan za potpunu konstitutivnu aktivnost Janus kinaze 2 u stanici.9 Postoje 2 teorijska modela mehanizma patogeneze mijeloproliferativnih poremeaja. U modelu A JAK2 (V617F) (sam ili uz joa jednu ili viae mutacija) je odgovoran za patogenezu onih pacijenata koji imaju mutaciju prisutnu, dok su druge mutacije odgovorne za patogenezu pacijenata bez mutacije JAK2 (V617F). U modelu B mijeloproliferativni fenotip se ve razvio kad se pojavila mutacija JAK2 (V617F). Pri tom mutacija samo poti e daljni razvitak bolesti. To bi zna ilo da pacijenti bez mutacije imaju rani stadij MPD-a. U oba modela prelazak iz heterozigotnog u homoziogotno stanje prestavja selektivnu prednost razvoju bolesti.7  Slika 1. Primjer signalnoga puta JAK2. Dimerizacija receptora prolaktinom, poti e fosforilaciju JAK2 koja zauzvrat fosforilira nizvodne signalne molekule STAT5. STAT 5 dimeriziraju i kao dimeri ulaze u jezgru kako bi potakli transkripciju ciljnih gena Mutacija JAK2 (V617F) u mijeloproliferativnim poremeajima U po etnim studijama je mutacija pronaena u granulocitima u 77 % pacijenata s PRV, u 35 % pacijenata s ET i u 43 % slu ajeva CIMF-e.8 No, daljnje studije koje su upotrebljavale osjetljivije metode detekcije izvjeauju da je ak 99 % PRV pacijenata, te oko 50 % CIMF pacijenata nosilo mutaciju.9 Iscrpno istra~ivanje na 806 slu aja ET pokazalo je da 53 % pacijenata ima mutaciju i da je u toj podpopulaciji u estaliji prijelaz iz ET u PRV.11 Mutacija u mijeloidnim stanicama mo~e biti heterozigotna i homozigotna. Homozigotnost je pronaena u etvrtini oboljelih PRV.3 Najviae je homozigota pronaeno u PRV pacijenata dok ih je u ET i CIMF bilo pacijenata mnogo manje. Ovakva raspodjela upuuje da postotak homozigotnosti direktno ovisi o postotku heterozigotnosti mutacije u populaciji pacijenata odreenog poremeaja.10 Mutacija JAK2 (V617F) je ste ena, ne postoje slu ajevi policitemija u kojima se unutar nekih obitelji ova mutacij javlja mnogo eae od slu ajnog. Tu se mo~e rei da se nasljeuje sklonost k razvijanju mutcije, ali mutacija nipoato nije nasljedna jer nije prisutna u spolnim stanicama.12 Uz klasi ne mijeloproliferativne poremeaje, mutacija JAK2 (V617F) se mo~e pronai i u drugim atipi nim MPD, poput hipereozinofilnog sindroma (1.4 % ) i mijelomonocitne leukemije (7.6 % ), ali i u rijetkim slu ajevima akutne mijeloi ne leukemije (AML) kojoj je prethodio neki mijeloproliferativni poremeaj, u 7.4 % slu aja bcr/abl negativne kroni ne mijeloi ne leukemije (CMML), u 2.6 % pacijenata s mijelodisplasti nim sindromom,2 i u ak 50 % pacijenata sa refraktornom anemijom, prstenastim sideroblastima i trombocitozom (RARS-T).12  SHAPE \* MERGEFORMAT  1.2.4 Utjecaj mutacije JAK2(V617F) i drugih imbenika na fenotip mijeloproliferativnih poremeaja ini se kako je mutacija JAK2(V617F) nu~na za razvijanje fenotipa klasi nih mijeloproliferacija. No, iako se mo~e pronai u veini slu ajeva mijeloproliferativnih poremeaja, o igledno nije jedini uzrok ovom fenotipu.10 Mutacija JAK2(V617F) je povezana sa veim komplikacijama u tijeku bolesti, poput sekundarne mijelofibroze, tromboze i krvarenja.7 Takoer,usmjerava multipotentne hematopoetske mati ne stanice razvitku u eritroidnu lozu.8 Homozigotnost dodatno poti e diferencijaciju u eritroidnu lozu, ali je ona manje pouz dan kriterij jer se javlja i u ET i CIMF pacijenata (iako mnogo rjee). Prijelaz iz heterozigotnosti u homozigotnost mogao bi objasniti i ne tako rijedak fenotipski prijelaz iz ET i rane PRV u kasnu PRV,te pojavljivanje sekundarne mijelofibroze koja prati kasnije stadije mijeloproliferativnih poremeaja.2 Homozigotnost je povezana i s duljim trajanjem bolesti.7 Pretpostavlja se da stanice koje imaju oba alela jak mutirana, imaju selektivnu prednost u proliferaciji i pre~ivljavanju naspram stanica koje su heterozigotne za mutaciju JAK2(V617F). Slika 3. Razvoj mijeloproliferativnih poremeaja. Mutacija V617F na jednom alelu gena jak2daje selektivnu prednost ovoj heterozigotnoj mutiranoj stanici nad normalnom stanicom. Gubitak heterozigotnosti (9pLOH) osigurava joa veu selektivnu prednost homozdfJNX"\"( (//==DDHI`ObOTTTTUZZ\\\BbFbddg"ghhiiijpqqDq2z6zJ{P{.|/|||}~Lƃ·Ї.4h B* mHnHphuh 5mHnHujh UmHnHu h 6 h 5 h H*h N   "$&(*,.>@$da$$$ d^$ `a$$$ d^$ `a$v??B@hJ\" (,./T//////356: $,d^,a$ $ & Fda$$da$ $d`a$:<=====B@lEGHHHIILbO>QQTTTTTRU $ d^$ ` $d`a$$da$RUTUnXZ\^FbiiiiiiiiijjlqqqBqDqrwd$da$$$ d^$ `a$w/|}~ & Fd$a$d$da$ $d`a$~4tz|8 $da$ $d`a$d4ntv’|28HLhntиָ tNP<^µܒh B*ph3fh CJaJh 6CJaJh 5CJaJUh 6CJaJmHnHuh CJaJmHnHuh CJaJ"jh CJUaJmHnHu h 5jh UmHnHujh U h H*h 3igotne mutirane stanice nad heterozigotnom mutiranom i normalnom stanicom. Na fenotip mutacije JAK2(V617F) mogu utjecati i druge mutacije,ato je i vjerojatno, jer su sve hematopoetske bolesti, osim CML (bcr/abl), posljedica viae od jedne mutacije. Inhibitori JAK2(V617F) kao potencijalni lijekovi mijeloproliferativnih poremeaja Mutacija JAK2(V617F) bi mogla biti inhibirana bilo kojim kinaznim inhibitorom airoke specifi nosti poput lestaurtiniba i erlotiniba, ili inhibitorom specifi nim za Janus kinaze 2. Velika veina ovih tvari inhibira i divlji tip Janus kinaze 2. Jaka inhibicija JAK2 uzrokovala bi kriti nu situaciju: anemiju, neutropeniju i trombocitemiju istovremeno.12 Stoga se od novih lijekova tra~i da inhibiraju JAK2(V617F) , ali ne i divlji tip JAK2. Obeavajua je terapija novom tvari ITF2357 koja inhibira proliferaciju stanica sa JAK2(V617F) mutacijom tako da smanji koli inu JAK V617F proteina i inhibira nizvodni signalni put . ITF2357 je u potpunosti uklonila fosforilirane JAK2 V617F, STAT5 i STAT3 ali nije utjecala na divlje tipove JAK2 i STAT proteina u kontrolnoj stani noj liniji.13 Osim ovih tvari, postoje i mnoge druge, koje, nakon uspjeanog laboratorijskog, ekaju na po etak klini kog testiranja. 2. CILJ RADA Analizirana je prisutnost mutacije JAK2(V617F) u DNA dobivenoj iz uzoraka krvi 45 pacijenata oboljelih od mijeloproliferativnih bolesti iz Klini kog bolni kog centra Split, te su postavljeni sljedei ciljevi istra~ivanja : Molekularnom analizom uzoraka DNA odrediti u estalost mutacije u pacijenata Odrediti raspodjelu po spolu,dobi i uputnoj dijagnozi Usporediti dobivene rezultate s podacima dostupnim iz literature 3. ISPITANICI I METODE Materijali Uzorci U ovom radu analiziran je 191 uzorak krvi pacijenata oboljelih od neke od mijeloproliferativnih bolesti. Uzorci su uzeti i molekularno analizirani u razdoblju od 15 mjeseci (20. rujna 2006. godine do 31. prosinca 2007. godine) u Laboratoriju za klini ku genetiku KBC Split. Puferi i otopine Tube za vaenje krvi Tube sa EDTA kao antikoagulansom za 3-5 mL krvi, Brand PLUS, Becton Dickinson Vacutainer Systems (Franklin Lakes, NJ, USA) Puferi i otopine koriatene za izolaciju DNA iz pune krvi Puferi/otopine sadr~ani u kompletu reagencija Puregene Genomic DNA Purification Kit, Gentra Systems (Minneapolis, MN, USA): Otopina za lizu eritrocita (od engl. Red Blood Cell Lysis Solution) Otopina za lizu leukocita (od engl. Cell Lysis Solution) Otopina za talo~enje proteina (od engl. Protein Precipitation Solution) Otopina za hidraciju DNA (od engl. DNA Hydration Solution) Izopropanol (2-propanol), 100%, Sigma (Highland, IL, USA) Etanol, 70%, Merck (Whitehouse Station, NJ, USA) Puferi, reagencije i enzimi koriateni u lan anoj reakciji polimeraze (PCR) za odreivanje JAK2 V617F mutacije PCR-pufer, 10x MgCl2, 25 mM Uzvodna uobi ajena po etnica, 10 M Nizvodna po etnica specifi na za mutaciju, 10 M Nizvodna kontrolna po etnica, 10 M Deoksinukeozid-trifosfati (dATP, dTTP, dCTP i dGTP), 2.5 M Voda, redestilirana AmpliTaq Gold"! DNA Polymerase, 5 U/L Sve reagencije su proizvod Applied Biosystems (Foster City, CA, USA). Puferi i otopine koriateni za elektroforezu u gelu agaroze TBE-pufer, 1X Tris 90 mM Borna kiselina 90 mM EDTA (pH=8) 2 mM Otopina je autoklavirana. Agarozni gel (1,5%) Agaroza, Electrophoresis grade, InvitrogenTM Life Technologies 1.5 g Etidij bromid 10 L TBE, 1X do 100 mL Agaroza je otopljena na cca 40C, a gel je (joa topao) rasporeen na prethodno pripremljene nosa e za gel. DNA marker, 50 bp DNA Ladder, InvitrogenTM Life Technologies Boja za uzorke, Blue Juice Gel DNA Loading Buffer, InvitrogenTM Life Technologies (Carlsbad, CA, USA) Laboratorijska oprema Centrifuge 5810R, Eppendorf AG, Hamburg, Njema ka Z 230 MR, Berthold Hermle AG, Gosheim, Njema ka PCR ureaj, GeneAmp PCR System 9700 Ureaj za elektroforezu, Consort E 844, 400 V  400 mA UV iluminator, 312 nm  254 nm Polaroid-kamera, FotoDyne 1-1440 (USA) Metode Izolacija leukocitne DNA iz pune krvi Uzorci krvi sa EDTA su do izolacije bili pohranjeni na 4C. Izolacija stani ne DNA vraena je Puregene Genomic DNA Purification Kit (Gentra Systems) komercijalnom kompletom reagencija i prema doti nom standardnom protokolu za izolaciju DNA iz 3 mL pune krvi (sadr~i eritrocite). Puna krv (1 volumen) se doda u Falcon tubu od 15 mL koja ve sadr~i 9 mL (3 volumena) otopine za lizu eritrocita (RBC Lysis Solution), promijeaa se i inkubira 5 minuta na sobnoj temperaturi. Prilikom inkubacije potrebno je joa minimalno jedanput izmijeaati uzorak. Centrifugiranjem (2 min, 2000 x g) istalo~e se itavi leukociti, supernatant se odlije, a tuba kratko osuai na upijajuem papiru. Talog se sna~no vrtlo~no promijeaa kako bi se stanice resuspendirale u ostatku tekuine. Na resuspendirane stanice se doda 3 mL (1 volumen) otopine za lizu leukocita (Cell Lysis Solution) i vrtlo~no se pomijeaa uzorak. Ako i dalje postoje nakupine stanica u uzorku, potrebno ga je inkubirati na 37C dok otopina ne postane homogena. Uzorak sa liziranim leukocitima se ohladi na sobnu temperaturu i doda se 1 mL (1/3 volumena) otopine za talo~enje proteina (Protein Precipitation Solution). Stani ni lizat se sna~no vrtlo~no promijeaa 20-ak sekundi i centrifugira se (5 min, 2000 x g) kako bi se stvorio vrsti smei talog proteina. Ako talog nije vrst, uzorak se ponovno vrtlo~no promijeaa, stavi na led 5 minuta i centrifugira. Supernatant koji sadr~i DNA se izlije u tubu sa 3 mL izopropanola i promijeaa se nje~nim preokretanjem tube dok se ne pojave bijele niti DNA koje stvaraju ~elatinoznu nakupinu. DNA se istalo~i centrifugiranjem (3 min, 2000 x g), odlije se supernatant i tuba kratko osuai na upijajuem papiru. Talog se ispire dodatkom 3 mL etanola i preokretanjem. Uzorak se kratko centrifugira (1 min, 2000 x g), etanol pa~ljivo odlije, a talog se osuai, prvo upijajuim papirom, a potom i na zraku 10-15 minuta. DNA se rehidrira u 250 L otopine za hidraciju DNA (DNA Hydration Solution) inkubacijom na 65C 1h i preko noi na sobnoj temperaturi. Dio se hidriranog uzorka kratko centrifugira i prenese u epruvetu za privremenu pohranu na 4C ili trajnu na -20C do -40C. Preostali se dio koristi za lana anu rekciju polimeraze kao DNA kalup. Umno~avanje egzona 12 gena JAK2 lan anom reakcijom polimeraze specifi nom za alel (allele-specific PCR) Egzon 12 je kodirajui dio JAK2 gena unutar kojeg nastaje G!T to kasta mutacija DNA koja rezultira V617F to kastom mutacijom JH2 regulatorne domene. Lan ana reakcija polimeraze je obavljena prema radu Baxter i sur., 2005.3 Upotrijebljene su dvije nizvodne po etnice ( forward primers ). Prva nizvodna po etnica je specifi na za mutirani alel i sadr~i namjerno nepoklapanje ( mismatch ) sa DNA kalupom divljeg tipa na treem nukleotidu od 3' kraja. Ovo nepoklapanje poboljaava specifi nost po etnice za mutirani alel i ona umna~a fragment DNA duga ak samo 203 bp. Druga nizvodna po etnica umna~a fragment duga ak 364 bp (i u divljem tipu alela i u mutiranom) i slu~i kao unutarnja kontrola za PCR.. Analiza umno~enih ulomaka agaroznom gel-elektroforezom Napravljena je horizontalna agarozna gel elektroforeza na 1,5 %-tnim gelom agaroze sa etidij-bromidom u 1X TBE-puferu. Svaki set uzoraka pratile su pozitivna kontrola (sigurno pozitivni DNA uzorak), negativna kontrola (PCR smjesa bez DNA kalupa) i komercijalni marker sa razmakom fragmenata od 50 bp. Na 5 L uzorka koriateno je 2 L plave boje za uzorke po jednoj ja~ici. Uvjeti elektroforeze su bili redom: napon 60 V, jakost struje 50 mA, trajanje 2h. Uzorci su promatrani na UV iluminatoru i rezultati zabilje~eni polaroid kamerom. 4. REZULTATI Analiza dobivenih podataka: Tablica 4. 1. Usporedba u estalosti mutacije JAK2 (V617F) pacijenata po spolu POZITIVAN NALAZNEGATIVAN NALAZSPOLU ESTALOSTUDIO (%)U ESTALOSTUDIO (%)MU`KARCI 827,61168,8}ENE2172,4531,6UKUPNO2964,41635,6  Slika 4.1. Usporedba u estalosti mutacije JAK2(V617F) po spolu Tablica 4.2. Usporedba u estalosti mutacije JAK2 (V617F) pacijenata po dobnim skupinama POZITIVAN NALAZNEGATIVAN NALAZSTAROST(god)U ESTALOSTUDIO (%)U ESTALOSTUDIO (%)0-4027,4422,241-5027,4316,751-60414,8527,861-70829,6527,8>701140,715,6UKUPNO2760,01840,0  Slika 4.2. U estalost mutacije JAK 2 (V617F) po dobnim skupinama Tablica 4.3. Usporedba u estalosti mutacije JAK2(V617F) u skupinama pacijenata po uputnim dijagnozama DIJAGNOZAPOZITIVAN NALAZNEGATIVAN NALAZU ESTALOSTUDIO (%)U ESTALOSTUDIO (%)PRV1965,51062,5ET26,916,25SyM620,7425CIMF26,916,25UKUPNO2964,41635,6  Slika 6.3. Usporedba u estalosti mutacije JAK (V617F) po uputnoj dijagnozi 5. RASPRAVA Mijeloproliferativne bolesti su klonalni poremeaji mati nih hematopoetskih stanica. Zajedni ko im je poja ano stvaranje i nakupljanje velikog broja nezrelih stanica mijeloidne loze u koatanoj sr~i, perifernoj krvi i u nekim tkivima i organima kao ato su jetra i slezena, ato dovodi do poveanja ovih organa. esto se zbog poveanog broja stanica u krvi mogu razviti mikrovaskularne smetnje, ali i ozbiljnije tromboze. Uzrok glavnim skupinama mijeloproliferativnih poremeaja  policitemiji rubra vera, esencijalnoj trombocitemiji i kroni noj idiopatskoj mijelofibrozi joa uvijek nije razjaanjen. Mutacija JAK2 (V617F) se nalazi u veini mijeloidnih stanica veine pacijenata. Ova mutacija najvjerojatnije daje neku selektivnu prednost mutiranoj stanici u odnosu na zdrave stanice i tako doprinosi patogenezi mijeloproliferativnih poremeaja. Za razliku od normalnih stanica, eritroidne mati ne stanice koje nose ovu mutaciju rastu u kulturi bez eritropoetina, pa se mo~e zaklju iti da mutacija JAK2(V617F) uzrokuje stani ni rasta i proliferacije neovisno od faktora rasta.3 Prema dobivenim rezultatima od ukupno analiziranih bolesnika (n=45) utvreno je da da je njih 29 bilo pozitivno na mutaciju JAK2 (V617F), a od toga je bilo pozitivna 21 ~ena (72,4%) i 8 muakaraca (27,6%). (Tablica 4.1; Slika 4.1.) Prema literaturi je poznato da je u estalost mutacije vea u starijim dobnim skupinama. Kako bi se utvrdilo postoji li povezanost mutacije s ~ivotnom dobi ispitivana je populacija podijeljena u aest dobnih skupina. Iz tablice 4.2 se vidi da je mutacija rijea do pedesete godine ~ivota (14,8%), a s porastom ~ivotne dobi, raste i udio mutacija (29%). Najvea u estalost mutacije je iza sedamdesete godine ~ivota (40,7%). Takoer je analizirana u estalost mutacije prema uputnoj dijagnozi.( Tablica 4. 3, Slika 4.3.). Po naaim podacima, mutacija je utvrena u gotovo 2/3 pacijenata (65.5%). Mutacija JAK2 (V617F) je esta u policitemiji rubra vera, a u literaturi njena u estalost varira od 77%-99%.9 U skupini bolesnika s ET i CIMF bilo je 6,9% pozitivnih nalaza, ali ih ne mo~emo uzeti u obzir zbog nedovoljnog broja ispitanika. Iscrpne studije izvjeauju da je ak 50% CIMF pacijenata i 53% ET pacijenata pozitivno na mutaciju JAK2 (V617F).11 Va~no je napomenuti da je u Laboratoriju za molekularnu genetiku na Zavodu za patologiju, sudsku medicinu i citologiju KBC Split ukupno analizirano 480 uzoraka (148 s pozitivnim nalazom i 332 s negativnim nalazom). Od tih 480 bilo je 205 muakaraca (42,7%) i 275 ~ena (57,3%). O toga broja, 72,4% ~ena i 27,6% muakaraca bilo je pozitivno na mutaciju JAK2 (V617F). Meutim, zbog nedostatnih ostalih klini kih podataka (godine, uputna dijagnoza) nije se mogla napraviti potpuna statisti ka analiza. Molekularna analiza mutacije JAK2 ima veliko zna enje u dijagnostici i lije enju hematoloakih bolesti, posebno mijeloproliferativnih poremeaja, te je za o ekivati da e sa veim brojem analiza postati joa va~nija u dijagnosticiranju bolesti, a veim brojem ispitanika i s potpunim kini kim podacima, dobili bi i vjerodostojnije rezultate. 6. ZAKLJU AK Na osnovu dobivenih rezultata analize DNA iz 45 uzoraka krvi na prisutnost mutacije JAK2 (V617F) metodom alelno specifi nog PCR-a mo~e se zaklju iti: Utvrena je u estalost pozitivnih nalaza u 29 od 45 analiziranih pacijenata. Od toga je bilo pozitivno 21 od 29 ~ena (72,4%), te 8 od 19 muakaraca (27,6%). Usporedbom u estalosti mutacije prema dobnim skupinama utvreno je da je u estalost mutacije rijea do pedesete godine ~ivota, a s porastom ~ivotne dobi raste i udio mutacija. Mutacija je naj eaa iza sedamdesete godine ~ivota (40.7%). Analizom prema uputnoj dijagnozi dokazana je najvea u estalost mutacije kod PRV (19/29; 65%), a kod ET i CIMF dokazana je mutacija u 6.9% ispitanika, ali zbog malog broja ispitanika uzorak nije reprezentativan. Kompletnom analizom DNA uzoraka na mutaciju JAK2 (V617F) obraeno je 480 ispitanika (148+, 332-), 215 muakaraca i 255 ~ena. Od toga je 29% ~ena i 20.1% muakaraca pozitivno na mutaciju Uzevai u obzir dobivene rezultate mo~e se zaklju iti da je pretraga na JAK2 (V617F) va~an dio dijagnosti kog algoritma mijeloproliferativnih poremeaja i da e potrebe za ovom pretragom rasti, dok bi klini ka vrijednost bila i vea kad bi imali potpune podatke. 7. SA}ETAK Policitemija rubra vera (PRV), esencijalna trombocitemija (ET), i kroni na idiopatska mijelofibroza (CIMF) su mijeloproliferativni poremeaji u kojima se nakupljaju krvne stanice mijeloidne loze. U ovim poremeajima primijeena je u estalost pojavljivanja mutacije V617F u autoregularnoj domeni Janus kinaze 2. Mutacija sprje ava normalnu inaktivaciju JAK2, te tako uzrokuje prekomjernu proliferaciju stanica. Rezultati analize uzoraka krvi koji su bili ispitivani na prisutnost mutacije JAK2 (V617F) metodom alelno-specifi nog PCR-a pokazuju da je mutaciju imalo 72.4 % ~ena i 27.6 % muakaraca. Mutacija se pojavljuje razli itom u estaloau meu pojedinim bolestima a najvea je kod PRV (65%). Usporedbom u estalosti mutacije po dobnim skupinama vidi se da je mutacija rijea do pedesete godine ~ivota a s porasti ~ivotne dobi raste i udio mutacija (40,7%). Pretraga na JAK2 (V617F) ima veliko zna enje u analizi hematoloakih bolesti, posebno mijeloproliferativnih poremeaja, a za o ekivati je da bi sa veim brojem analiza i potpunim klini kim podacima joa dobila na vrijednosti. 8. SUMMARY Polycytemia rubra vera (PRV), essential thrombocytemia (ET) and chronic idiopathic myelofibrosis (CIMF) are myeloproliferative disorders in which myeloid blood cells accumulate. In this disorders there is a frequent appearance of mutation V617F in auto regulatory domain of Janus kinase 2. The mutation prevents normal inactivation of JAK2, thereby enhancing proliferation of cells. The results of blood samples analysis that were closely examined for mutation JAK2 (V617F) by allele-specific PCR, indicated that 72.4 % of women and 26.6% had that mutation. The mutation occurs in various incidences between specific illnesses, but it is most common in PRV (65%) In comparing the incidence with age, it seems that mutation is seldom before the age of fifty. However, incidence of this mutation increases proportionally with ageing, especially after 70 years (40.7%). Testing on JAK2 (V617F) has a strong impact on analysis of haematological diseases,especially myeloproliferative disorders. Hopefully, this type of testing with complete clinical data could additionally contribute to accurate diagnostics. 9. LITERATURA Labar B, Hauptmann E i sur. Hematologija. `kolska knjiga, Zagreb, 1998 Michiels JJ, Bernema Z, Van Bockstaele D, De Raeve H, Schroyens W. Current diagnostic criteria for the chronic myeloproliferative disorders essential thrombocytemia, polycytemia vera and chronic idiopathic myelofibrosis. Pathologie Biologie 2007; 55; 92 - 104 Baxter EJ, Scott LM, Campbell PJ, East C, Fourouclas N, Swanton S, Vassiliou GS, Bench AJ, Boyd EM, Curtin N, Scott MA, Erber WN, Green AR. Acquired mutation of the tyrosine kinase JAK2 in human myeloproliferative disorders. Lancet 2005; 365; 1054- 1061 Chiat Y, Condat B, Cazals  Hatem D, Rufat P, Atmani S, Chaoui D, Guilmin F, Kiladjian JJ, Plessier A, Denninger MH, Casadevall N, Valla D, Briere JB. Relevance of the criteria communly used to diagnose myeloproliferative disorders in pacients with splanchic vein trombosis. Br J Haematol 2005; 129: 553  60 Vrhovac B, Franceti I, Jakai B, Labar B, Vuceli B. Interna medicina. Naklada Ljevak, Zagreb, 2003 Zhao R, Xing S, Li Z, Fu X, Li Q, Krantz SB, Zhao ZJ. Identification of an acquired JAK2 mutation in polycytemia vera. J Biol Chem 2005; 280: 22788  92. Kralovics R, Guan Y, Prchal T. Acquired uniparental disomy of chromosome 9p is a frequent stem cell defect in polycytemia vera. Exp Hematol 2002; 30:229  236. James C, Ugo V, Casadevall N, Constantinescu SN, Vainchenker WA. JAK2 mutation in myeloproliferative disorders: pathogenesis and therapeutic and scientific prospects. Trends Mol Med; 2005; 11 (12): 546  54. Levine RL, Gilliand DG. JAK2 mutations and their relevance to myeloproliferative disease. Curre Opin Hematol 2007; 14: 43  7. Ihle JN, Gilliand DG. Jak2: Normal function and role in hematopoietic disorders. Curr Opin Genetics Dev 2007; 17: 8  14. Campbell P, Scott LM, Buck G. Definition of subtypes of essential trombocytemia and relation to polycytemia vera based on JAK2 V617F mutation status: A prospective study. Lancet; 2005: 366 (9501): 1945  53. Skoda R. The genetic basis of myeloproliferative disorders. Hematology 2007; 1: 1  10. Guerini V, Barbui V, Spinelli O, Salvi A, Dellacasa C, Carobbio A, Introna M, Barbui T, Goalay J, Rambaldi A. The histone deacetylase inhibitor ITF2357 selectively targets cells bearing mutated JAK2(V671F). Leukemija 2008; 22 (4): 740  7. 10. }IVOTOPIS Ime i prezime: Tea Kutla a Datum i mjesto roenja: 20. sije nja 1985. god., Split, Hrvatska Obrazovanje: Osnovna akola: Ravne njive  Split Srednja akola: Zdravstvena akola  Split, smjer medicinska sestra/tehni ar, 1999-2003 Medicinski fakultet Sveu iliata u Splitu, 2003- 2009 Studentska razmjena: Finska 2008. Strani jezik: aktivno znanje engleskog jezika     PAGE  PAGE 43 Slika 2 Mehanizam aktivacije JAK2. A) u inaktivnom stanju JAK2 je vezana za receptor bez citokina. Pseudokinazna (JH2) domena inhibira kinaznu (JH1) domenu. B) Vezanje citokina za receptor uzrokuje strukturne promjene u receptoru i JAK2. JH2 otpuata JH1 domenu. JAK2 se aktivira. 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