ࡱ>  []RSTUVWXYZM 4bjbj== .WW^l 8 $ ,< "B B B    R:T:T:T:T:T:T:$9= Y?x:     x:1B B =<111 F$B B R:1 R:1"17:R:B  `x c1":R:S<0<&:,@14@R:1PREGLEDNI LANAK / REVIEW PAPER ONKOGENI I PROTOONKOGENI  pregled aktualnih istra~ivanja Oncogenes and protooncogenes  review of recent researches Marko Hohateter1, Maja Popovi2, }eljko Grabarevi1 1 Zavod za opu patologiju i patoloaku morfologiju 2Zavod za biologiju Veterinarski fakultet Sveu iliata u Zagrebu SA}ETAK Onkogeni su geni koji poti u autonomni stani ni rast u tumorskim stanicama. Karakterizira ih sposobnost da poti u stani ni rast bez prisustva normalnih mitogenih signala. Protoonkogeni su geni koji su sastavni dio genoma svake stanice i oni su fizioloaki regulatori stani ne proliferacije i diferencijacije. Proteini koje kodiraju onkogeni se nazivaju onkoproteini i sli ni su normalnim produktima protoonkogena tj. protoonkoproteinima, osim ato onkoproteini ne sadr~e elemente va~ne za kontrolu regulacije stani nog ciklusa. Na taj na in aktivnost onkogena u transformiranim stanicama postaje trajna i nije ovisna o faktorima rasta ili drugim vanjskim signalima koji utje u na stani ni. U ovom lanku su ukratko opisane dosadaanje spoznaje o onkogenima i protoonkogenima, te su opisana neka aktualna istra~ivanja na ovom podru ju, koja su raena na svinji, kao pokusnom modelu, koji je jedan od naj eae istra~ivanih u veterinarskoj medicini. KLJU NE RIJE I: onkogen, protoonkogen, svinja, tumor ABSTRACT Oncogenes are genes which promotes cell growth in tumors cells. They are characterized by ability to promote cell growth without presence of normal mithogen signals. Protooncogenes are genes which are constitutive parts of cell genom and works as physiological regulators of cell proliferation and differentiation (Kumar at al., 2005). Proteins which are encoded by oncogenes are oncoproteins and they are very similar to normal products of protooncogenes or to the protooncoproteins, with exception that oncoproteins do not have elements which are important to cell cycle regulation. Because of that, oncogens are permanently active in the transformed cells and they are not dependent of growth factors or others signals which have influence on cell cycle. In this paper recent achievements about oncogenes and protooncogenes are shortly described. We have described a different familyes of protooncogenes and oncogenes, mechanisms of their activity and their influence to development of tumors. We have also described some new data on this field which were conducted in pigs as a experimental model which is one of the most often animal used in veterinary medicine. KEY WORDS: oncogene, protooncogene, pig, tumor ONKOGENI Onkogeni su geni koji upravljanju kontrolom rasta i diferencijacijom neoplasti nih stanica. Do prvih spoznaja o postojanju takvih gena doalo se istra~ivanjem retrovirusa. Tipi ni retrovirus ima genom koji se sastoji do 3 gena: gag  kodira kapsidne proteine; pol  kodira reverznu transkriptazu; env  kodira proteine ovojnice (Slauson, 2002). Takve gene imaju neki sporotransformirajui virusi (Virus miaje leukemije, Virus pti je leukoze, Virus leukemije ma aka) (Slauson, 2002). Ti su virusi infektivni i oni izazivaju nastanak limfoma i leukemija u organizmu domaina. Njih karakterizira dugi inkubacijski period potreban da bi se razvila neoplasti na bolest. Osim njih postoje i akutno-transformirajui retrovirusi koji induciraju stvaranje neoplazmi u vrlo kratkom vremenu. Ti virusi nisu jako infektivni i izolirani su iz razli itih tumora npr. Virus Rousov-og sarkoma kod pilia (Kumar i sur, 2005). Takvi virusi uz tri prethodno nabrojana gena sadr~e i src gen. Taj gen je potreban za transformaciju stanice i naziva se virusni onkogen (v-onc). Dosad je poznato oko 20 virusnih onkogena povezanih s akutno-transformirajuim retrovirusima. Ti virusi imaju skraeni gen, a skraeni dio je zamijenjen sa v-onc. Takvi se virusi mogu replicirati samo u stanicama koje su konificirane sa tzv. pomonim kompetentnim virusom koji nadomjeata funkcije retrovirusa koje mu nedostaju. PROTOONKOGENI Spoznaje o virusnim onkogenima dovele su do otkria da stanice u organizmu kod svih ~ivotinja sadr~e homologne sekvence genoma koji se, kako bi se naglasila njihova sposobnost transformacije nazivaju protoonkogeni. Protoonkogeni su visoko konzervirani ubikvitarni geni koji se nalaze u svih organizama na evolucijskoj ljestvici (Kumar i sur., 2005). Danas se zna da su onkogeni opisani u akutno-transformirajuih retrovirusa u stvari protoonkogeni uklopljeni u virusni genom i promijenjeni u virusne onkogene. Struktura i funkcija protoonkogena u stanici organizma mo~e se pod djelovanjem kemijskih i dr. kancerogena promijeniti na taj na in da dobbiju sposobnost transformacije stanica. Na taj na in promijenjen protoonkogeni se nazivaju stani ni onkogeni (c-onc) (Slauson, 2002). Dosad je otkriveno oko 100 razli itih protoonkogena. Funkcija protoonkogena je kodiranje proteina koji imaju ulogu u regulaciji stani nog ciklusa i diferencijaciji stanice (Grabarevi, 2002). Postoji znatna kvalitativna i kvantitativna razlika izmeu regulacijskih proteina za rast i transformaciju stanice kodiranih izmeu protoonkogena i od strane stani nih ili virusnih onkogena. Produkti mnogih onkogena su funkcionalni analozi proteina uklju enih u normalne signalne putove u stanici koji zapo inju vezanjem liganda za receptor na stani noj membrani, a zavraavaju stani nom proliferacijom ili promjenom u nekoj drugoj stani noj funkciji (Grabarevi, 2002). VRSTE ONKOGENA I MEHANIZAM DJELOVANJA Onkogeni iji su proteinski produkti locirani u jezgri imaju ulogu u regulaciji transkripcije i ekspresiji gena. Tu spadaju myc (c-myc, n-myc, l-myc); c-myb i c-fos onkogeni (Slauson, 2002). Oni se ve~u na DNK i reguliraju transkripciju mnogih stani nih gena. Produkti myc i fos onkogena su takoer povezani sa malim ribonukleoproteinima iz stani ne jezgre koji sudjeluju u posttranskripcijskoj transformaciji RNK. Oba onkogena i myc i fos su eksprimirani tijekom prelaska stanice iz Go u G1 fazu stani nog ciklusa, te su va~ni za ekspresiju drugih gena va~nih u mitoti ki aktivnih stanica. Veina proteinskih produkata onkogena pokazuje svoje djelovanje vezanjem na receptore na stani noj membrani ili u citoplazmi, te su to proteini uglavnom s protein-kinaznom aktivnoau ili proteini koji kataliziraju fosforilaciju tirozinskih skupina ili treonin-serinskih skupina. Na taj na in fosforilacija proteina predstavlja najva~niji mehanizam u regulaciji stani nih funkcija. Ta grupa onkogena ulazi u src - gensku porodicu (Slauson, 2002) i oni svi imaju homolognu tirozin-kinaznu sekvencu. Neki od tih onkogena kodiraju i faktore rasta, pa je tako genski produkt sis onkogena faktor rasta podrijetlom iz trombociota (platelate derived growth factor) koji je sna~an mitogen za fibroblaste, endotelne i dr. stanice. ras porodicu onkogena ine tri gena: H ras, otkriven kod Virusa Harvey-evog miajeg sarkoma; K-ras, otkiriven kod Virusa Kirsten-ovog miajeg sarkoma i N-ras, stani ni onkogen izoliran iz stanica neuroblastoma (Slauson, 2002). Ti su geni visoko evolucijski konzervirani te su naeni i kod ni~ih organizama npr. kod kvasaca. Proteini kodirani od ras -onkogena su blisko vezani sa G proteinima koji se nalaze na citoplazmatskoj strani stani ne membrane i imaju ulogu u prijenosu signala od stani nih receptora na citoplazmatske sekundarne glasnike. bcl-2 porodica onkogena je jedna od najkasnije otkrivenih skupina onkogena i njihovi produkti su smjeateni na membrani mitohondrija, a funkcija im je u regulaciji apoptoze tj. programirane stani ne smrti. Oni zajedno sa produktima bax  onkogena odreuju da li e stanica ui u apoptozu ili pre~ivjeti i na taj na in uzrokovati proliferaciju tkiva. `to je joa jedan od dokaza da su onkogeni povezani sa neoplasti nom transformacijom.  Slika 1. Stani na lokalizacija i funkcija najzna ajnijih skupina protoonkogena. Protoonkogeni su obojeni crvenom bojom. (Kumar i sur., 2005) Figure 1. Cellular localisation and function of major clases of protooncogenes. The protooncogenes are colored red. (Kumar at al., 2005)  Slika 2. Prikaz aktivnosti ras protoonkogena. Mutirani ras (onkogen) je trajno aktiviran zbog nemogunosti da hidrolizira GTP, ato dovodi do trajne mitoti ke stimulacije stanice bez prisustva vanjskog signala. (Kumar i sur., 2005) Figure 2. Model of action of ras protooncogene. The mutant ras (oncogene) is permanently activated because of inability to hydrolyze GTP, leading to continous stimulation of cells without any external trigger. (Kumar et al., 2005) ULOGA ONKOGENA U RAZVOJU TUMORA Postoje brojni podaci koji upuuju na to da je zna ajna uloga onkogena pri nastanku tumora (Kumar i sur., 2005). Glavni dokaz za to dolazi od injenice da su akutno transformirajui virusi sposobni in vitro transformirati stanice, a in vivo uzrokuju stvaranje tumora. Fibroblasti transformirani djelovanjem virusa Rous-ovog sarcoma su tumorogeni ni kada se inokuliraju u pilie. Specifi na uloga v - src onkogena u tom je slu aju potvrena na taj na in da mutacija tog gena u obliku delecije rezultira gubitkom transformacijske sposobnosti virusa, dok je klonirana v - src DNK sposobna transformirati stanice u koje je unesena (Slauson, 2002). Vrlo brzi razvoj tumora koji je karakteristi an za akutno-transformirajue viruse je u jakoj suprotnosti sa sporim razvojem veine spontano nastalih tumora. Usprkos tome postoji viae dokaza da onkogeni imaju va~nu ulogu u razvoju spontano nastalih tumora. Transfekcijski test u kojem se DNK iz tumorskih stani nih linija ili tumorskog tkiva unese u stanice glodavaca pokazuje da 15 do 20% humanih tumora ima aktivne onkogene sposobne za transformaciju stanica domaina (Slauson, 2002). Veina od njih su onkogeni iz ras porodice onkogena. myc  onkogeni su takoer jako nazoni kod nekih neoplazmi, i to 20 do 80 puta viae nego kod zdravih stanica. Kod Burkitt-ovog je limfoma translokacija myc - onkogena na imunoglobulinski lokus vezana uz u estalost limfoidnih neoplazmi. I u mnogim drugim slu ajevima mutacije protoonkogena imaju va~nu ulogu u stvaranju transformacijske sposobnosti. ras - onkogeni nastaju to kastim mutacijama pa se njihovi proteini mijenjaju u odnosu na fizioloake samo za pojedinu aminokiselinu (Slauson, 2002). Oni imaju sekvence homologne s G proteinima koji imaju sposobnost vezanja i hidrolize GTP-a. Dok se normalni G proteini aktiviraju vezanjem GTP-a, a inaktiviraju njegovom hidrolizom, kod ras onkoproteina dolazi do blokade hidrolize pa je na taj na in ras G protein kontinuirano aktivan ato dovodi do aktivacije stani nih funkcija koje utje u na stani ni rast i diferencijaciju. Dokazano je da su ras  onkogeni aktivni u stanicama tumora mlije ne ~lijezde, karcinoma ko~e, timoma, tumora bubrega i papiloma ko~e. Osim to kastih mutacija mogue su i druge strukturne promjene koje dovode do aktivacije onkogena. Kod kroni nih mijeloi nih leukemija u ljudi dolazi do translokacije gena izmeu kromosoma 9 i 22 ato rezultira stvaranjem hibridnog gena koji kodira fuzijski protein sastavljen od v-abl onkogena i v erb-B onkogena (Slauson, 2002). V-abl onkogen kodira tirozin-kinazu ija je aktivnost jako poveana kod fuzijskog proteina, dok v-erb-B onkogen kodira dio receptora za epidermalni faktor rasta, bez njegove ekstracelularne kontrolne domene, pa je zato tirozin-kinazna aktivnost kontinuirana. Danas se smatra da osim kod akutno-transformirajuih retrovirusnih onkogena nije dovoljna aktivacija samo jednog onkogena da bi doalo do razvoja tumora (Kumar i sur., 2005). Dokaz za to je ato ras-onkogen mo~e transformirati fibroblaste iz stani nih linija no ne i primarne embrionalne fibroblaste. Smatra se da je uzrok tome to ato su stanice iz linijskih kultura ve proale neke genetske promjene koje pridonose njihovoj besmrtnosti. myc-onkogen takoer nije samostalno dostatan za tumorogenezu ve je potrebna aktivnost i dodatnog onkogena. Infekcija stanica sa virusom koji sadr~i samo myc onkogen dovodi do hiperplasti nih promjena u tkivu, dok infekcija s virusom koji sadr~i samo ras-onkogen dovodi do displasti nih promjena. Ako doe do infekcije sa virusom koji nosi oba i ras- i myc-onkogen dolazi do stvaranja malignih neoplazmi. Iz svih ovih iznesenih podataka vidljiva je sposobnost onkogena, osobito ako ih viae zajedni ki djeluje, za indukciju neoplasti nih bolesti. PREGLED AKTUALNIH ISTRA}IVANJA ONKOGENA I PROTOONKOGENA NA SVINJI KAO POKUSNOM MODELU U ovom poglavlju opisana su neka aktualna istra~ivanja onkogena i protoonkogena koja su raena na svinji kao pokusnom modelu. U literaturi se tumori kod svinja relativno rijetko opisuju. Smatra se da je to posljedica toga ato veina svinja ne do~ivi starost potrebnu da bi se razvio tumor (`imec i sur., 2001). Meu opisanim neoplasti nim bolestima kod svinja su naj eaa akutna diferencirana mijelocitna leukemija (Grabarevi, 2002), tumori jajnika (hemangiomi, maligni luteomi i metastatski adenokarcinomi) (`imec i sur., 2001), a tumor koji se relativno eae javlja u svinja je kutani melanom. Taj tumor se naj eae javlja u Sinclairove minijaturne svinje, pa ta ~ivotinja slu~i kao animalni model za prou avanje kutanog melanoma (Grabarevi i sur., 2005). Iako je iz navedenih podataka vidljivo da svinje rijetko oboljevaju od tumora, one su kao pokusni modeli, zbog svoje fizioloake sli nosti sa ovjekom, ~ivotinje koje se naj eae koriste u istra~ivanjima koja se provode na podru ju onkogena i protoonkogena. Naj eae su istra~ivani protoonkogeni i onkogeni iz porodica myc-, fos- i jun-onkogena. Istra~ivana je uloga svinjskog s-myc protoonkogena koji kodira nuklearni fosfoprotein koji ima va~nu ulogu u mnogim fizioloakim procesima (Reiner i sur., 1999). Najva~niji procesi u kojima on sudjeluje su stani na proliferacija, diferencijacija, apoptoza; hormonska regulacija; regulacija faktora rasta, citokina, limfokina; u razvoju i diferencijaciji; u adipogenezi, miogenezi i dr. c-myc protoonkogen funkcionira kao slo~eni molekularni prekida  koji na mnogobrojne signalne podra~aje iz stanice reagira ili pozitivo ili negativno i na taj na in regulira mnogobrojne procese u stanici. Strukturalna analiza je pokazala da je c-myc visoko evolucijski konzervirani gen koji je 90-100% homologan u svih ~ivotinjskih vrsta. Kod ljudi je taj gen smjeaten na HSA8 kromosomu dok je kod svinja utvreno da se on nalazi na SSC 4p13 kromosomskoj poziciji. U istra~ivanju je izoliran c-myc iz svinjskih hepatocita. Lan anom reakcijom polimeraze je utvreno da se sastoji od tri egzonska dijela i tri promotorska mjesta, a kodira protein sastavljen od 439 aminkoiselina. S obzirom da je poznato da je c-myc protoonkogen uklju en u adipogenezu istra~ivanja tog gena su va~na radi utvrivanja njegovog mogueg utjecaja na varijabilnost u nakupljanju masti u svinja, ato bi se moglo iskoristit kod dobivanja komercijalnih pasmina svinja. Takoer je istra~ivana ekspresija svinjskih n-myc onkogena u razvoju bubrega i onkogenezi (Benedit i sur., 1991). Utvrena je ekspresija n-myc RNK u stanicama bubrega u novoroene prasadi do starosti od 5 tjedana. U tome razdoblju zavraava glomerularna diferencijacija, a imunohistokemijski je utvreno da je n-myc RNK selektivno eksprimiran u zametnim epitelnim stanicama bubrega. Izdvajanjem tih stanica i uzgojem u kulturi stanica te koinjekcija mutiranog ras onkogena i n-myc u te stanice dovela je do stvaranja ~arianih stani nih nakupina u tekuem agaru, gubitka kontaktne inhibicije i stvaranjem trajne stani ne linije. Iz navedenih je spoznaja vidljivo da n-myc gen ima va~nu ulogu u razvoju bubrega, te njegova povezanost kao jednog od imbenika povezanog sa neoplasti nom transformacijom. c-myc protoonkogen je prou avan i u granuloza stanicama svinje tijekom razli itih faza rasta jajnog folikula (Sato i sur., 1994). Analizirana je c-myc mRNK izolirana iz granuloza stanica metodom northern blot hibridizacije. Utvrena ja znatna koli ina c-myc mRNK u granuloza stanicama malih folikula (1-2 mm) dok u srednjim (3-5 mm) i velikim (6-11 mm) nije utvrena prisutnost c-myc transkripta. Utvreni transkript veli ine je 2,4 kb i po svojoj je veli ini jednak c-myc mRNK izoliranom iz humanih granuloza stanica. Opisani rezultati upuuju da ekspresija c-myc protoonkogena u ranom stadiju rasta folikula mo~e imati ulogu u atutonomnom stani nom rastu nezrelih granuloza stanica i u ranoj folikulogenezi ato upuuje na to da bi ta faza razvoja folikula mogla biti neovisna o hipofizno gonadotropnoj stimulaciji. Na ekspresiju protoonkogena mogu utjecati razni egzogeni agensi, pa je tako prou avan utjecaj nefrotoksina diklorvinilcisteina (DCVC) na ekspresiju c-fos i c-myc onkogena u kulturi stanica svinjskog bubrega komparativno sa dijelovanjem razli itih faktora rasta i 12-O-tetradekanoilforbolacetata (Vamavakas i Koster, 1993). DCVC izaziva direktno oateenje DNK i kidanje veza na dvostrukoj uzvojnici DNK. Dodavanjem 0,2% DCVC u inkubacijski medij nakon 24 sata je doalo jasno vidljive ekspresije c-fos i c-myc u stanicama iz kulture. Sli ne razine ekspresije c-fos mRNK su inducirane kombinacijom faktora rasta (EGF, inzulin, transferin), a ekspresija c-myc mRNK sa 12-O-tetradekanoilforbolacetatom. No kod DCVC je razina ekspresije rasla za vrijeme cijelog perioda inkubacije za razliku faktora rasta i 12-O-tetradekanoilforbolacetata kada su ekspresijske razine rasle od 0,5 do 1 sat poslije po etka inkubacije, a nakon toga dolazi do njihovog pada. Provedena je analiza ekspresije c-fos u endometriju nazimica tijekom spolnog ciklusa i za vrijeme ranog graviditeta (Dubois i sur., 1993). Utvreno je da je ekspresija c-fos m RNK jednaka tijekom spolnog ciklus i ranog graviditeta (do 18. dana) u svim danima osim 12. dana kada je ekspresija c-fos znatno vea kod gravidnih nazimica. Takoer je u tom razdoblju kod obje skupine lokaliziran imunoreaktivni fos protein u ~lijezdanim stanicama, povrainskim epitelnim stanicama i u stanicama miometrija. Rezultati ove studije upuuju na to da je porast endometrijske ekspresije c-fos mRNK povezan sa razvojem maternalnog prepoznavanja graviditeta u nazimica. Istra~ivana je regulacija c-fos, c-jun, jun-B i c-myc protoonkogena u stani noj kulturi svinjskih Leydig-ovih stanica pod djelovanjem gonadotropina (hCG) i razli itih faktora rasta (EGF, FGF, transformirajui faktor rasta beta, inzulinu sli ni faktor rasta I) (Hall i sur., 1991). Iz dobivenih je rezultata vidljivo da cCG dovodi do porasta ekspresije c-fos (do 9 puta), jun-B (do 18 puta) i c-myc (do 5 puta) , ali ne utje e na ekspresiju c-jun.U inak EGF i FGF na c-fos, i jun-B je slabiji nego kod cCG, ali je ekspresija c-myc mRNK ja a. Takoer se utvrdilo da da EGF i FGF potenciraju u inak hCG na c-fos i jun-B, dok hCG potencira u inak faktora rasta na c-jun. Transformirajui faktor rasta beta poja ava samo ekspresiju jun-B, a inzuliu sli ni faktor rasta I pove ava ekspresiju c-fos, jun-B i c-myc, ali slabije od hCG. Iz svih je ovih podataka vidljivo da je regulacija ekspresije istra~ivanih protoonkogena u Leydig-ovih stanica multifaktorijalna. Vidljivo je i da postoji razli ita reakcija lanova porodice jun protoonkogena na odreene faktore rasta. Ovi rezultati podr~avaju hipotezu da fos i jun protoonkogenski proteini imaju ulogu u dugotrajnoj regulaciji odgovora na razli ite hormone i faktore rasta kojima su Leydigove stanice izlo~ene in vivo. Raena su i istra~ivanja o ekspresiji protoonkogena u stanicama miokarda svinje vezano za ishemiju i reperfuziju tkiva (Brand i sur., 1992). Molekularna osnova adaptacije miokarda na ishemiju i reperfuziju je slabo poznata. Misli se da jezgrini protoonkogeni dijeluju kao tercijarni prenosioci koji prevode citoplazmatski signal u dugotrajne promjene u ekspresiji gena. Raena je studija o ekspresiji protoonkogena EGR-1, c-fos, fos-B, c-jun, jun-B i c-myc u stanicama miokarda pod utjecajem ishemije i reperfuzije. Protoonkogeni su bili razli ito eksprimirani. Nije zabilje~ena ekspresija fos-B protoonkogena. c-myc gen je bio eksprimiran, ali razina mu se nije mijenjala ovisno o ishemiji. Ekspresija c-jun je bila malo poja ana dijelovanjem ishemije dok su c-fos, EGR-1 i jun-B bili znatno inducirani (5 do 7 puta). Ovo istra~ivanje je pokazalo da u adaptivnom odgovoru miokarda na ishemi ni stres ulogu imaju najmanje 4 protoonkogena tj. transkripcijska faktora koja mogu imati ulogu u reparacijskim procesima i angiogenezi. Osim razli itih kemijskih agenasa na ekspresiju protoonkogena utjecaj mogu imati i razli ita zra enja, ato je joa jedan od dokaza indukcije tumora zbog aktini kog dijelovanja . Martin i suradnici su istra~ivali indukciju c-fos u usporedbi sa c-jun protooonkogena tijekom rane reakcije ko~e svinja na gama zrake (Martin i sur., 1993). Oba protoonkogena su istovremeno inducirana, kod visokih doza (8 do 48 GY). Ja ina indukcije je i vremenski i o dozi ovisna, ali je ja a c-fos ekspresija. Niske doze radijacije (0,5 do 2 Gy) ne induciraju c-jun, dok je c-fos znatno induciran. Kao ato je ve prethodno opisano postoje brojne sli nosti izmeu pojedinih virusnih gena i odreenih stani nih protoonkogena pa se u tom smjeru istra~ivao genom virusa afri ke svinjske kuge i sli nost njegovih gena sa protoonkogenom bcl-2 i genom Epstein-Barr virusa BHRF1 (Neilan i sur., 1993). Utvreno je da se dio genoma virusa afri ke svinjske kuge LMW5-HL izoliran iz svinjskih makrofaga pokazuje visoku sli nost sa protoonkogenom bcl-2, a u manjoj mjeri s BHRF1 genom Epstein-Barr virusa. Ova sli nost upuuje na moguu ulogu bcl-2 protoonkogena u pre~ivljavanju stanica tijekom infekcije s virusom afri ke svinjske kuge. Iz iznijetih je podataka vidljivo da je uloga protoonkogena u ~ivotu stanica, a posebice u regulaciji stani nog ciklusa i stani noj diferencijaciji vrlo va~na, te da postoje brojna nerijeaena pitanja vezana uz same mehanizme njihovog djelovanja. Takoer ova istra~ivanja nedvojbeno potvruju klju nu ulogu onkogena kod razvoja neoplasti nih promjena. LITERATURA: Benedit I, Rich MA, Moldwin R, Waber p, Steinberg BM, Nisen PD. N-myc oncogene expression in porcine renal development and oncogenesis. Pediatr Res 1991; 29 (3); 268-271. Brand T, Sharma HS, Fleishmann KE i sur. Proto-oncogene expression in porcine myocardium subjected to ischemia and reperfusion. Circ Res 1992; 71 (6); 1351-1360. Dubois DH, Simmen RC, Fliss A, Smith LC, Bazer FW. Expression of c-fos in porcine endometrium during the estrous cycle and early pregnancy. Biol Reprod 1993; 49 (5); 943-950. Grabarevi }. Veterinarska onkologija. Zagreb: DSK-Falco,2002. Hall SH, Berthelon MC, Avallet O, Saez JM. Regulatoon of c-fos, c-jun, jun-B, and c-myc messenger ribonucleic acids by gonadotropin and growth factors in cultured pig Leydig cell. Endocrinology 1991; 129 (3); 1243-1249. Kumar V, Abbas AK, Fausto N. Neoplasia. U: Robbins and Cotran Pathologic basis of disease. 7th edition . Philadelphia: Elsevier Saunders, 2005. Martin M, Piton P, Crechet F, Lefaix JL, Daburon F. Preferential induction of c-fos versus c-jun protooncogene during the immediate early response of pig skin to gamma-rays. Cancer Res 1993; 53 (14); 3246-3249. Neilaan JG, Lu Z, Afonso CL, Kutish GF, Sussman MD, Rock DL. An African swine fever virus gene with similarity to the proto-oncogene bcl-2 and the Epstein-Barr virus gene BHRF1. J Virol 1993; 67 (7); 4391-4394. Reiner G, Hecht W, Leeb T, Brenig B, Robic A, Dzapo V. Isolation and characterisation of the porcine c-myc proto-oncogene and chromosomal assignment to SSC 4p13. Animal Genetics 1999; 30 (3); 204-206 Sato A, Bo M, Maruo T, Yoshida S, Mochizuki M. Stage-specific expression of c-myc messenger ribonucleic acid and porcine granulosa cells erly in follicular growth. Eur J Endocrinol 1994; 131 (3); 319-322. Slauson DO, Cooper BJ. Mechanisms of dissease a textbook of comparative general pathology - 3th edition. St. Louis, Mosby a Harcourt Health Sciences Company, 2002. `imec Z, Grabarevi }, Artukovi B i sur. Ovarian hemangioma in the wild boar (Sus scrofa) A case report. Z. Jagdwiss 2001; 47; 268-274. Vamvakas S, Koster U. The nephrotoxin dichlorovinylcysteine induced expression of the protooncogenes c-fos and c-myc in LLC-PK1 cells- a comparative investigation with growth factors and 12-O-tetradecanoylphorbolacetate. Cell Biol Toxicol 1993; 9 (1); 1-13. PAGE  PAGE 8 &>@.0NPln~:@`j%%++++++,,,,,,$.*.0.6...2233 55T5^55566@677L9V9; ;<<<ڼjEHUmHsHCJEHmHsH6EH]mHsH EHmHsH5CJEH"\CJEHmHsHCJEH 5EH\EHI@B.0& dfhj|5% $`a$$a$$a$$a$3 2"F%t*v*x**h/ 5L9<<<<==>>>>>>>>>?$`a$$a$<<==????:?@?r?x?@@AAJAPA FF\GjG LL:L@LlMxMNNPPR"R:SDSTV^VxVVVV|WWFZNZ0\4\b]h]&^.^^^^^z`$aiiiii jjllnnppXrbrtt@uJuvvww^yhyEH 5EH\6EH]mHsHjMEHUmHsH EHmHsH5EH\mHsHR??@@BBBBBBBBBBJ&TXZ_v`x`z`&a$bgii.tlz`$`a$$a$hylzvz{{d|n|`}j}~~~~΀Ą(2|hnČƍ(DNnxΐؐ v &0X` HʚКښ Μ؜@X^h"BL6EH]mHsHEH 6EH] EHmHsH[lz΀<\HZ«/ޭ~S#&`#$ & F$a$$`a$ģΣʥԥbl",!"#%&,-./0340JmHnHu0J j0JU EHmHsH6EH]mHsHEH 6EH]#$%01234 & F&`#$h]h,1h. 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