ࡱ> 9DN56789:;<=>?@ABCQtv x z | ~ BDFHJLNPRez~`vbjbjt%4h~z -Fz"ۛ< BBBBBBB$)HhJ.B!BgE222B2B22Tn @g$!*^<`GE0-FKNKTKT`7:q2Q777BBڄX777-F$ SVEU ILI`TE U ZAGREBU VETERINARSKI FAKULTET Nikica Prvanovi, dr. vet. med. Uloga trofoblast proteina i leptina u ranoj dijagnostici gravidnosti i ranoj embrionalnoj smrtnosti u goveda Disertacija Zagreb, 2006. Doktorska disertacija izraena je na Klinici za porodniatvo i reprodukciju Veterinarskog fakulteta Sveu iliata u Zagrebu Predstojnik: prof. dr. sc. Antun Tomaakovi Voditelj: prof. dr. sc. Antun Tomaakovi POVJERENSTVO ZA OCJENU DISERTACIJE: prof. dr. sc. Marijan Cergolj dr.sc. Tihomira Gojmerac (znanstveni savjetnik, HVI) prof. dr. sc. Antun Tomaakovi Mojoj obitelji& & Zahvale Zahvaljujem se mentoru prof. dr. sc. Antunu Tomaakoviu kao i prof. dr. sc. Marijanu Cergolju na pru~enim savjetima i podraci, zatim mr. sc. Predragu Ko ili i Jurju Grizelju na pomoi prilikom izvoenja prakti nog dijela rada. Jednako tako zahvaljujem prof. dr. sc. Jean-Franoisu Beckersu, dr. sc. Jos Sulonu i dr. sc. Josipu Reaetiu na provoenju laboratorijskih testova i interpretaciji rezultata. Popis koritenih kratica: BCS-body condition score BPL-bovine placental lactogene BTP- bovine trophoblast proteins BPSP- bovine pregnancy specific proteins ET-embriotransfer ELISA- enzyme linked imuno assay ECG- equine chorionic gonadotropin HCG- human chorionic gonadotropin IDO-indoleamin 2,3 hidroksioksidaza IFN- interferon tau NT-nuklearni transfer, transfer jezgre NK stanice-natural killer stanice OTP- ovine trophoblast proteins OtR-oksitocinski receptor P4-progesteron 4 PAG- pregnancy associated glycoproteins PSP60-pregnancy specific protein 60 PGF2(-prostaglandin 2 alfa RIA- radio imuno assay UO-umjetno osjemenjivanje Sadr~aj: Uvod (cilj i svrha istra~ivanja)& & & & & & & & & & & & & & & & ..& ..7 PREGLED LITERATURE:& & & & & & & & & & & & & & & & & & ..& ...9 2.1. Rana embriogeneza i maj insko prepoznavanje gravidnosti& & & & ....9 2.2. Mehanizam placentacije i endokrinologija gravidnosti u pre~iva a ......14 2.2.1.Morfoloake, histoloake i fizioloake odlike placente i placentacija& .14 2.2.2.Transport preko placente tijekom gravidnosti& & & & & & & & & ..17 2.2.3. Placenta kao endokrini organ tijekom gravidnosti& & & & & & & ..19 2.3 Specifi nosti endokrinologije gravidnosti u goveda& & & & & & & & ....21 2.4 Laboratorijska, klini ka i ultrazvu na dijagnostika rane gravidnosti u pre~iva a& & & & & & & & & & & & & & & & & & & & & & & & & & & & & .& .22 2.5 Uloga trofoblast proteina (PAG) u maj inskom prepoznavanju gravidnosti i odr~avanju gravidnosti u goveda& & & & & & & & & & & & & & & & & & & 25 2.5.1. Blokiranje maj inog imunosnog odgovora tokom gravidnosti: implikacije za imunosupresiju& & & & & & & & & & & & & & & & & & & & & & & & & & & & 25 Uloga placente i binuklearnih stanica u razvoju i odr~avanju rane gravidnosti u goveda& & & & & & & & & & & & & & & & & & & & ..29 2.6. Primjena trofoblast proteina (PAG) u dijagnosti ke svrhe& & & ...31 2.6.1.Odreivanje trofoblast proteina (PAG, PSPB i PS60) pomou RIA & 31 Primjena trofoblast proteina (PAG) u dijagnostici gravidnosti & & & 33 2.6.3.Praenje gravidnosti i dijagnostika rane embrionalne smrtnosti s pomou PAG profila & & & & & & & & & & & & & & & & & & & & & & & & & & & ..35 2.7.Biokemijska srodnost leptina i ostalih citokina vezanih za gravidnost (pregnancy associated cytokines)& & & & & & & & & & & & & & & & .& & 37 2.8.Uloga leptina u reprodukciji & & & & & & & & & & & & & & & & ..39 2.8.1. Uloga leptina u odr~anju reproduktivne sposobnosti& & & & & & & . 39 Utjecaj leptina na spolno dozrijevanje& & & & & & & & & & .& .& & .41 Uloga leptina u gravidnosti& & & & & & & & & & & & & & & & & .& .41 Utjecaj leptina na funkciju jajnika& & & & & & & & & & & & & ..& & ..43 2.8.5. Povezanost razine leptina i laktacije& & & & & & & & & & & .& & & .43 3. MATERIJAL I METODE& & & & & & & & & & & & & & & & & & & & ...45 3.1.Odabir plotkinja& & & & & & & & & & & & & & & & & & & & & & & & ...45 3.2. Uzimanje krvi i ultrazvu no praenje steonosti & & & & & & & & & & ..47 3.3. Odreivanje progesterona, leptina i PAG u serumima primjenom RIA& & ..49 3.3.1.Osnovni principi seroloake reakcije RIA& & & & & & & & & & & & & & & 49 3.3.2 Odreivanje progesterona u serumu& & & & & & & & & & & & & & & & .51. 3.3.3. Odreivanje leptina u serumu& & & & & & & & & & & ..& & & & & & & ..52 3.3.4.Odreivanje PAG u serumu& & & & & & & & & & & & & & & & & & & & 54 4. REZULTATI ISTRA}IVANJA& & & & & & & & & & & & & & & & & & .57 4.1.Tabli ni prikaz rezultata za skupinu gravidnih krava& & & & & & & & & 57 4.2.Tabli ni prikaz rezultata za skupinu krava s embrionalnom smrtnosti& ..58 4.3.Tabli ni prikaz rezultata za skupinu negravidnih krava& & & & & & & & 59 4.4.Usporedni grafi ki prikaz leptina, PAG i progesterona u svih krava u Istra~ivanju& & & & & & & & & & & & & & & & & & & & & & & & & & & 67 4.5 Statisti ka analiza dobivenih vrijednosti& & & & & & & & ..70 4.5.1.Deskriptivna statistika za skupinu gravidnih krava 70 Deskriptivna statistika za skupinu krava s embrionalnom smrtnosti..71 Deskriptivna statistika za skupinu negravidnih krava...72 Analiza varijance i testiranje srednje vrijednosti koncentracije leptina, PAG-trofoblast proteina i progesterona izmeu pojedinih skupina krava..73 Faktori linearne korelacije istra~ivanih parametara& & & & & ............80 5.RASPRAVA& & & & & & & & & & & & & & & & & & & & & & & & & & .84 6.ZAKLJU CI& & & & & & & & & & & & & & & & & & & & & & & & & & .89 7.SA}ETAK& & & & & & & & & & & & & & & & & & & & & & & & & & & .90 8.ABSTRACT& & & & & & & & & & & & & & & & & & & & & & & & & & ..91 9.POPIS LITERATURE& & & & & & & & & & & & & & & & & & & & & .....92 10.}IVOTOPIS & & & & & & & & & & & & & & & & & & & & & & & & & .108 1.UVOD Odreivanje interferona i ostalih trofoblast proteina (PAG;BPSP) sve se viae primjenjuje kao objektivni dokaz postojanja ~ivog vitalnog embrija, a provodi se seroloakom metodom RIA. Naime, migracija blastociste u maternici, prije nidacije, redovito je popraena tvorbom ve navedenih proteina, interferona i ostalih citokina, ato ima za posljedicu imunosupresiju te osiguravanje optimalnih uvjeta za nidaciju. Oni nadalje djeluju antiluteoliti ki tako da sprije e dostatnu sintezu, otpuatanje i transport PGF2alfa do jajnika. Poti u tvorbu PGE na atetu PGF2alfa, a kod svinja atoviae pretvaraju sekreciju PGF2alfa iz endokrine (u lumen krvnih ~ila) u egzokrinu (u lumen maternice), ime onemoguuju kontakt luteolizina s ~utim tijelom. Cjelokupna se pojava naziva maj insko prepoznavanje trudnoe ili "maternal recognition of pregnancy". Osobito ju je va~no detektirati stoga jer je na taj na in mogue dokazati da je graviditet postojao, ak i ako je nakon toga nastupila rana embrionalna smrtnost, ato nam omoguuje bolje poznavanje a samim tim i suzbijanje navedene patologije te kona no omoguuje kvalitetnu provjeru uspjeha svih postupaka asistirane reprodukcije. Naime dosada se rana dijagnostika gravidnosti provodila dokazivanjem progesterona u krvi i mlijeku, ato ne dokazuje prisutnost embrija ve samo ~utog tijela, te klini kim i ultrazvu nim pregledom. Mana svih navedenih metoda je u tome ato ih je mogue provesti samo na ~ivotinji i nije mogue naknadno (iz seruma) dokazati da je u majci postojao ~ivi plod.. Odreivanjem trofoblast proteina mogue je objektivno dokazati gravidnost ve 24. dan. Metoda je primjenjiva kod svih vrsta domaih ~ivotinja, a osobito je dobro prou ena u pre~iva a. Pritom su uo ene razlike izmeu pojedinih vrsta domaih ~ivotinja pa se tako npr. bovini trofoblast proteini, za razliku od ov jih, izlu uju i nakon 38. dana gravidnosti. Unakri~ne reakcije izmeu ov jeg interferona-tau i goveeg trofoblast proteina dokazuju da je rije  o sli nim ali ne i identi nim molekulama. U posljednjih nekoliko godina dokazivanje trofoblast proteina provodi se u krava, ovaca, koza i svinja, a neato je slabije istra~eno u kuja i kobila. Zbog svoje neosporne va~nosti za daljnji napredak svih dosad primjenjivanih biotehnoloakih postupaka interferoni i ostali trofoblast proteini se danas vrlo intenzivno istra~uju te postoje tendencije da postanu rutinski dijagnosti ki postupak. U novije vrijeme sve je o itija i uloga leptina ija koncentracija direktno utje e na tijek i ishod gravidnosti, pojavu prvog estrusa po porodu i sli no. Leptin je hormon podrijetlom iz adipocita, koji se prete~no sintetizira u masnom tkivu odakle se lu i u opu cirkulaciju u omjeru koji je upravo proporcionalan koli ini masnog tkiva. U fizioloakim koncentracijama (2-10 ng/ml serum) on poti e steroidogenezu jajnika i razvoj embrija. Naprotiv vrlo visoka razina leptina (oko 100 ng/ml seruma) djeluje inhibitorno. Koncentracija leptina zna ajno raste u ranoj gravidnosti (u prva dva tjedna) i smatra se da regulira invazibilnost citotrofoblasta tijekom implantacije i placentacije. Leptin djeluje na embrij tako ato poveava unos i metabolizam glukoze putem regulacije metabolizma i sinteze inzulina. Sve navedeno je dokazano kod ljudi, miaeva, atakora i ovaca. Pokusi na ovcama su pokazali da porast koncentracije leptina u gravidnih ovaca potje e ispo etka od citotrofoblasta, potom od placente te kona no od masnog tkiva ploda. Koncentracija leptina u krvi ostaje poviena tijekom cijele gravidnosti i rapidno opada nakon poroda kod ljudi, ovaca, mieva i takora. Leptin je po svojoj strukturi biokemijski vrlo srodan placentalnom laktogenu i interferonu-tau, to ukazuje na mogunost da bi oni mogli imati i sli nu funkciju. Navedeno nas je dovelo na ideju da kod goveda na osnovi usporedbe koncentracija leptina i PAG tijekom rane gravidnosti pobli~e istra~imo mehanizam rane embrionalne smrtnosti te pre~ivljavanja embrija prije i u tijeku implantacije, a na osnovi komparacije kontrolne negravidne skupine, gravidne skupine kod koje je doalo do rane embrionalne smrtnosti i gravidne skupine krava. Svrha i cilj istra~ivanja O ekujemo da emo na temelju rezultata provedenog istra~ivanja retrogradno uo iti pojavu rane embrionalne smrtnosti u uzgojima krava obuhvaenim istra~ivanjem te da emo biti u prilici provjeriti nalaze dobivene ginekoloakom i ultrazvu nom pretragom te uspjeanost primijenjenog umjetnog osjemenjivanja i embriotransfera. Sve zajedno bi trebalo pomoi u razjaanjenju uzroka navedene patologije i njenom pravovremenom uklanjanju te posljedi no poboljaati plodnost mlije nih goveda na navedenim gospodarstvima. 2.PREGLED LITERATURE 2.1 Rana embriogeneza i maj insko prepoznavanje gravidnosti Gravidnost u goveda i razvoj ploda sastoji se od itavog niza faza koje su popraene odgovarajuim nazivljem. Nakon singamije (stapanja muake i ~enske jezgre u inu oplodnje) zigota se pretvara u embrij. Embrij je rana faza razvoja koju ne mo~emo razlikovati unutar razli itih vrsta tj. nemogue je primjerice razlikovati svinjski od goveeg embrija.Nakon placentacije embrij prelazi u fetus. Fetus je potencijalni potomak koji ve posjeduje odlike vrste kojoj pripada (ovca, krava, konj, slon, ovjek ili neato tree) a joa uvijek se nalazi u maternici (Senger, 2003). Zametak je posljedica za ea a koristi se kao zajedni ki naziv za embrij i fetus. Da bi uope doalo do placentacije i embrij postao fetus neophodna su 4 predstadija (Senger, 2003): razvoj unutar zone pellucide izlazak blastociste iz zone pellucide maj insko prepoznavanje gravidnosti tvorba ekstraembrionalnih ovojnica Neposredno nakon oplodnje uo avamo veliku stanicu tzv. Ootidu u kojoj su joa vidljive muaka i ~enska jezgra. To je jedna od najveih stanica u organizmu koju karakterizira relativno velika masa citoplazme u usporedbi s jezgrom. Ona je potrebna kako bi poslu~ila kao izvor citoplazme za brojne male stanice koje e uskoro nastati diobom. Ootida nakon stapanja muake i ~enske jezgre postaje zigota koja potom podlije~e itavom nizu diobi tzv. brazdanju. Stanice koje nastaju brazdanjem nazivaju se blastomere. Svaka blastomera posjeduje potencijal da se razvije u zdravog i kompletnog potomka. Jednojaj ani blizanci nastaju kao posljedica odvajanja blastomera. itav fenomen naziva se totipotencija a za 2-,4- i 8- stani ne blastomere ka~emo da su totipotentne. Pokusi u laboratoriju su pokazali i dokazali da je to mogue. Naime, izdvajanjem blastomera i njihovim prebacivanjem u surogat zonu pellucidu a potom embriotransferom u uterus primateljice iste vrste doalo je do normalnog razvoja ploda i posljedi no raanja ~ivog i vitalnog potomka kod kunia, konja, goveda i ovaca (Senger, 2003). `toviae pokusi na ovcama u kojima je u anuklearne oocite umetnuta jezgra somatskih stanica odraslih ovaca ato je imalo za posljedicu normalan razvoj i raanje ~ivog i vitalnog janjeta iz ega se mo~e zaklju iti kako sve tjelesne stanice mogu biti potencijalno totipotentne, naravno pod odreenim uvjetima. Unutar zone pellucide stanice se dijele odnosno brazdaju sve dok ne dosegnu stadij morule. Za vrijeme razvoja unutar zone pellucide broj stanica se mijenja ali im je volumen stalan jer je limitiran zonom pellucidom. Morulu karakterizira dioba stanica na vanjski kompaktniji i unutraanji sloj stanica. Vanjski sloj stanica se vrsto pove~e, a natrij-kalij pumpa upumpava ione u meustani ne prostore. Potom u njih osmozom prodire voda i u moruli se akumulira tekuina koja stvara pritisak na vanjske stanice koje se spljoate a izmeu vanjskog i unutraanjeg sloja stanica se stvara aupljina koju nazivamo blastocoele. Unutraanji sloj stanica se polarizira i tvori unutraanju stani nu masu, dok vanjski spljoateni sloj stanica tvori trofoblast. Potom slijedi izlazak iz zone pellucide ili hatching odnosno valjenje. Valjenje se odvija pod djelovanjem 3 sile (Senger, 2003): rast i nakupljanje tekuine unutar blastociste sinteza enzima unutar trofoblasta kontrakcije blastociste U svakom slu aju blastocista se sastoji od unutraanje stani ne mase (Senger, 2003) iz koje e se razviti plod i trofoblasta iz kojeg e se razviti horion, fetalna komponenta placente odnosno posteljice. Diobe u blastocisti potpoma~u nakupljanje tekuine i porast unutraanjeg tlaka. Jednako tako, trofoblast lu i proteoliti ke enzime koji slabe i razgrauju zonu pellucidu. Potom se sama blastocista po inje kontrahirati i relaksirati ato izaziva nagle promjene unutraanjeg tlaka i dodatno poti e pucanje i raspad zone pellucide. Kada nastane mala pukotina na zoni pellucidi, blastocista se provu e kroz nju i potom se oslobaa u lumenu maternice. Da bi u tom mediju mogla pre~ivjeti neophodna je aktivnost ~utog tijela jajnika, adekvatna sinteza progesterona u ~utom tijelu i zadovoljavajui odgovor sluznice maternice na djelovanje progesterona. Nakon valjenja dolazi i do naglog rasta blastociste i poveavanja ukupne mase. Tako primjerice govei embrij starosti 13 dana dose~e duljinu od 3 mm a 17-tog dana zajedno s trofoblastom dose~e duljinu od 250 mm i dolazi do fenomena kojeg nazivamo elongacija. Sa 18 dana embrij se izdu~i tako da se prote~e kroz oba roga. Tako postignuti kontakt sa sluznicom maternice slu~i maj inskom prepoznavanju gravidnosti. Joa br~i rast embrija zabilje~en je kod svinja iji embriji nakon valjenja rastu brzinom od 4-8 mm na sat (Barb i sur. 2001). Paralelno s rastom embrija, intenzivno se razvijaju i plodne ovojnice: ~umanj ana vreica, horion, amnion i alantois. Sve su podrijetlom iz trofoblasta, endoderma, mezoderma i samog embrija. Za sve papkare, pa tako i goveda karakteristi an je filamentozni izgled navedenih ovojnica, dok kod konja itav zametak s ovojnicama zadr~ava sferi nu strukturu. Trofoblast zajedno sa primitivnim endodermom i mezodermom daje osnovu iz koje izrasta horion i amnion. Horion se potom ve~e za endometrij a amnion se pretvara u tekuinom ispunjenu vreu koja hidrauli ki atiti plod od mehani kih povreda. }umanj ana vrea se pak razvija iz primitivnog endoderma. Ona je samo krae vrijeme prisutna kao plodna ovojnica i nestaje s daljnjim razvojem embrija, ali upravo ona daje zametnu osnovu za genitalni tuberkulum. Nakon ato se horion razvije u strukturu koja podsjea na krila i obavije embrij, po inje se formirati amnion. Amnionska tekuina tokom razvoja atiti tkiva embrija od meusobne adhezije. Amnionski mjehuri kod goveda je palpabilan ve sa 30-45 dana gravidnosti kao maleni mjehuri izra~enog turgora. Kako je embrij u ovoj fazi izrazito osjetljiv, palpaciju je potrebno izvoditi izrazito pa~ljivo. U toj fazi se po inje razvijati i alantois koji nastaje iz evaginacije primitivnog crijeva, a njegova je funkcija da skuplja tekui ekskret ploda.Kako se horion i alantois razvijaju i rastu tako se njihova kontaktna povraina poveava ili ak fuzira tvorei alantohorion koji predstavlja fetalnu komponentu placente. Paralelno s rastom i razvojem zametak je i biokemijski aktivan kako bi omoguio svoje pre~ivljavanje u maternici i izazvao maj insko prepoznavanje gravidnosti ija je svrha spre avanje imunoloakog odbacivanja i ubijanja zametka te spre avanje luteolize. Naime za pre~iva e je karakteristi no da sintetiziraju oksitocin koji stimulira endometrij na otpuatanje prostaglandina koji izazivaju lutelizu. U krava i ovaca blastocista sintetizira tvari koje blokiraju sintezu materni nih oksitocinskih receptora. To su specifi ni proteini koji spadaju u razred interferona. Interferoni su nespecifi ni glikoproteini koje sintetiziraju leukociti, fibroblasti, limfociti i trofoblast. Oni imaju antivirusnu aktivnost i moduliraju aktivnost ciljanih tkiva. Relativno su male molekularne mase (18,000 do 20,000 daltona) a ukoliko je dolo do koncepcije prisutni su u endometriju 13-21 dan nakon ovulacije. Oni ne izazivaju porast koncentracije progesterona dakle ne djeluju luteotropno, ve se ve~u na endometrij i blokiraju sintezu oksitocinskih receptora. Takoer se ve~u i za apikalni dio materni nih ~lijezda i poti u sintezu i otpuatanje proteina klju nih za pre~ivljavanje embrija. Zanimljivo je da interferon (tau) ne djeluje na porast koncentracije progesterona u serumu tj. nema luteotropno djelovanje. U krma a embrij otpuata estradiol koji preusmjerava sekreciju PGF2( iz endokrine (u krvne ~ile) u egzokrinu (u materni ni lumen) (Arthur i sur., 2001.) Naime, estradiol djeluje na sintezu prolaktinskih receptora a prolaktin regulira transport kalcija kroz membrane i posljedi no utje e na transport svih molekula pa tako i na preusmjeravanje PGF2( koji nakon ato je izlu en u lumen viae nema na ina da se vrati u cirkulaciju i izazove luteolizu. Estradiol kojeg lu e svinjski embriji takoer je va~an i stoga ato poti e kontrakcije miometrija ija je funkcija da pomognu migracije plodova i njihov adekvatan i proporcionalan smeataj u oba roga. Svinjski embriji takoer lu e interferon tau ali njegova funkcija nije do danas razjaanjena. Prili no je sigurno da nema istu funkciju kao kod pre~iva a (Arthur i sur., 2001). Jednako tako, pokazalo se da je neophodno da krma a u svakom rogu ima barem po dva zametka. Naime, ukoliko krma a nema dovoljno zametaka, uslijediti e luteoliza i zameci e biti resorbirani. Kod kobile je migracija ploda kroz maternicu joa izra~enija nego kod svinja jer je uniparna vrsta. `toviae, pokusi su pokazali da je migracija ploda preko cijele povraine rogova presudna za maj insko prepoznavanje gravidnosti u kobila. Da bi se graviditet odr~ao konjski embrij mora migrirati 12-14 puta dnevno i to tijekom 12., 13. i 14. dana gravidnosti kako bi blokirao luteolizu. Migracija embrija je neophodna jer je konjski embrij sferi an, za razliku od embrija u pre~iva a (krava, ovca, koza, koauta) koji kontakt sa endometrijem posti~u posredstvom elongacije. U svakom slu aju i konjski embrij sintetizira i tvori specifi ne proteine ija je funkcija zasada nepoznata. Ljudski embrij pak lu i hCG (humani chorionski gonadotropin) ija je funkcija inhibicija intraovarijske luteolize. Lu i ga embrij 7-9 dana nakon luteolize. Sam mehanizam djelovanja nije joa poznat. Najmanje se joa zna o maj inskom prepoznavanju gravidnosti u kuja i ma aka. Do sada nije izoliran protein specifi an za embrije pasa i ma aka i smatra se da mo~da i ne postoji te da nije potreban za pre~ivljavanje embrija. Naime ~uto tijelo kod kuje ne ovisi o gravidnosti a njegov nastanak, tijek i regresija su identi ni kod gravidnih i negravidnih kuja nakon estrusa. Kod ma aka kojima je ovulacija inducirana koitusom ili uope ne doe do stvaranja ~utog tijela ve folikul koji nije ovulirao regresira i ciklus se ponavlja za 8-10 dana jer ovulacije nije niti bilo ili pak dolazi do parenja i ovulacije, a ~uto tijelo koje posljedi no nastaje ne ovisi o nastanku, izostanku ili tijeku gravidnosti. Ono ato je definitivno zajedni ko svim vrstama domaih sisavaca to je da svi trebaju relativno visoku konstantnu razinu progesterona u krvi kako bi se graviditet odr~ao premda su mehanizmi koji ju omoguavaju kao i izvori progesterona tokom gravidnosti (~uto tijelo, placenta) bitno razli iti i direktno ovise o pojedinoj vrsti (Senger, 2003; Arthur i sur., 2001) Slika 1: Antitluteoliti ki mehanizam djelovanja interferona- tijekom rane gravidnosti-bazira se na blokiranju sinteze PGF2( preko oksitocinskih receptora  Slika 2: A i B Interakcije embrija i endometrija u goveda  2.2.Mehanizam placentacije i endokrinologija gravidnosti u pre~iva a 2.2.1.Morfoloake, histoloake i fizioloake odlike placente i placentacija Placenta ili posteljica predstavlja privremeni organ koji nastaje u ranoj gravidnosti i posjeduje biokemijsku i hormonalnu aktivnost tijekom intrauterinog ~ivota ploda. Ona je mjesto metaboli ke mjene izmeu zametka i majke, ali je i mnogo viae od toga. Osim ato predstavlja mjesto komunikacije izmeu krvotoka majke i ploda ona ima i itav niz endokrinih funkcija a to su (Senger, 2003): -odr~avanje gravidnosti -stimulacija mlije ne ~lijezde majke -poticanje rasta ploda U humanoj medicini mo~emo govoriti o implantaciji kao fenomenu koji prethodi i zapo inje placentaciju. Zapravo je rije  o uranjanju embrija u sluznicu maternice za koju se ve~e i kao takva je prisutna samo kod primata i glodavaca. Kod domaih kopitara, papkara i meso~dera nemamo takvu pojavu ve se embrij jednostavno ugnijezdi (pove~e, nidira) sa sluznicom endometrija i placentacija mo~e po eti. Placenta se sastoji od horionskih resica ploda te od modificiranih slojeva maternice ija je redukcija vrsno specifi na i predstavlja tip prilagodbe optimalan za rast i razvoj zametka ovisno o vrsti kojoj pripada. Openito placenta predstavlja mjesto metaboli ke izmjene cirkulacije ploda i majke, te predstavlja privremeni endokrini regulatorni organ. Endokrina uloga placente je dvojaka; tijekom gravidnosti osigurava odr~anje gravidnosti a pred termin poroda inicira po etak poroda. Endokrinoloaka regulacija poroda vrlo je slo~ena jer osim izbacivanja ploda i plodnih ovojnica porod inicira itav niz promjena u organizmu plotkinje; po etak i odr~anje laktacije, involuciju maternice i restituciju endometrija te povratak cikli ne aktivnosti jajnika. Od ukupno 3 plodne ovojnice-alantoisa, amniona i horiona, samo horion tvori placentu. Funkcionalna jedinica placente je horionska resica. Horionske resice su sitne prstolike izbo ine na povraini horiona koje uranjaju u endometrij i spajaju se s njim. Na osnovi razmjeataja i dubine uranjanja resica u endometrij (broj slojeva koje dijele cirkulaciju majke i ploda tj. stupanj redukcije endometrija) razlikujemo razli ite tipove placente koje nalazimo u domaih sisavaca. Na osnovi razmjeataja horionskih resica placenta domaih sisavaca mo~e biti anatomski gledano (Okljeaa, 1956): Placenta difusa (kobila, krma a) Placenta zonaria (meso~deri) Placenta discoidalis (primati, glodavci) Placenta cotiledonaria (pre~iva i) Placenta u kobile spada u skupinu difusa ato zna i da su horionske resice rasporeene po itavoj povraini placente ali istovremeno ima mnoatvo malenih mikrozona s resicama pa mo~emo rei da je rije  o mikrokotiledonima. Sli nu pojavu nalazimo i na placentama u krma e. Placentu kobile karakterizira i pojava endometrijskih aaica koje su takoer endokrinoloaki aktivne i lu e eCG. One su prisutne na maternici od 35 do 60 dana gravidnosti. U kobile se plod ve~e sa sluznicu 24. dana, a sama placentacija uslijedi do 40. dana nakon oplodnje. Placentu zonariu kakvu imaju meso~deri karakterizira centralna prstenasta zona na kojoj se nalaze horionske resice. Oko nje se nalazi pigmentirana zona koju sa injavaju sitni krvni ugruaci. Tu pigmentiranu zonu nazivamo joa i paraplacenta a smatra se da ima va~nu ulogu u prijenosu ~eljeza s majke na plodove. Najudaljenija od centralne zone je prozirna transparentna zona koja je slabo vaskularizirana a ija je funkcija vjerojatno vezana uz direktnu apsorpciju iz lumena maternice. Placenta diskoidalis koju imaju primati i glodavci redovito tvori jedan ili dva diska koji su jedina mjesta s horionskim resicama. Placenta kotiledonaria koju imaju pre~iva i, posjeduje tvorbe sli ne butonima koje nazivamo kotiledoni, a njihova veli ina, oblik i razmjeataj su vrsno specifi ni. Tako ovce i koze imaju 90-100 placentoma konkavnog oblika, dok krave, koaute, srne i ~irafe imaju 70-120 konveksna placentoma (oblika gljive) i na osnovi veli ine placentoma mo~emo procijeniti i pribli~nu gestacijsku starost ploda. Placentomi se sastoje od karunkula koji potje u od placente materne i kotiledona koji potje u od placente fetalis.Kod ovce se ve 16. dana nakon oplodnje diferenciraju karunkuli na sluznici maternice dok se kod krave oni pojavljuju 25. dana nakon oplodnje. Prije toga je placenta u nastajanju zapravo placenta diffusa. Nakon toga dolazi do procesa placentacije koji zavraava stvaranjem kona nog spoja kod ovce ve 30. dana, a kod krave 40. dana nakon oplodnje. Tokom samog fetalnog rasta i razvoja se placentomi viaestruko poveaju ovisno o opsegu mjene tvari izmeu ploda i majke i kod krave pred porod imaju i po 5-6 cm u promjeru. Placentu mo~emo odrediti i prema broju slojeva koje posjeduje odnosno prema stupnju redukcije endometrija u svrhu lakae izmjene tvari. Naime, placenta se sastoji od dvije krvne ~ile od kojih jedna pripada plodu a druga majci, kako je poznato histoloaki se svaka krvna ~ila sastoji od 3 sloja: endotela ili intime krvne ~ile, potom miainice i kona no vezivnotkivne rahle adventicije. Drugim rije ima ako su obje krvne ~ile intaktne placenta bi se trebala sastojati od 6 slojeva. Pojedine vrste sisavaca su pak u svrhu olakaavanja izmjene tvari izmeu majke i ploda reducirale pojedine slojeve placente materne tj. endometrija pa tako histoloaki mo~emo podijeliti placente u 4 osnovna tipa (Senger, 2003.): 1.epiteliochorialis (kobila, krma a)-6 slojeva 2.syndesmochorialis (pre~iva i)-5 slojeva 3.endoteliochorialis (meso~deri)-4 sloja 4.haemochorialis (primati, glodavci)-3 sloja Osobito je zanimljivo napomenuti da placentu domaih i divljih pre~iva a karakteriziraju divovske binuklearne stanice. Naime, one su podrijetlom iz trofoblasta a migriraju u placentu maternu tvorei pritom stani ni sincicij i fuziraju se u trinuklearne stanice. Prilikom fuzije otpuataju sadr~aj citoplazmatskih granula koje sadr~e BPSP (bovine pregnancy specific proteins). BPSP nazivamo joa i bPAG (bovine pregnancy associated glycoproteins). Divovske binuklearne stanice se javljaju kod ovaca 14. dana nakon koncepcije, dok se kod krave javljaju 18.-20. dana. One se stvaraju kontinuirano tijekom gravidnosti u pre~iva a a ine oko 20% svih stanica placente. Tijekom razvoja, binuklearne stanice migriraju iz epitela horiona i invadiraju epitel endometrija. Funkcija divovskih binuklearnih stanica je transport slo~enih molekula iz placente fetalis u placentu maternu. Nadalje, dokazano je da uz ve spomenuti BPSP (bovine pregnancy specific protein), ove stanice tvore i lu e i placentalni laktogen. Jednako tako u njima se odvija intenzivna steroidogeneza pri emu nastaju progesteron i estrogen. 2.2.2. TRANSPORT PREKO PLACENTE TIJEKOM GRAVIDNOSTI: Izmjena tvari izmeu cirkulacije ploda i majke svakako je prva i osnovna zadaa placente. Pri tom se ona odvija preko tri razli ita mehanizma: jednostavna difuzija, potpomognuta difuzija i aktivni transport. Plinovi i voda prolaze placentalnu barijeru na osnovi razlike koncentracija-jednostavna difuzija. Placenta sadr~i pumpe za aktivni transport kalcija, natrija i kalija. Glukoza i druge slo~enije molekule poput npr. aminokiselina prelaze placentalnu barijeru pomou potpomognute difuzije, vezane na specifi ne nosa e. Glukoza je osobito bitna jer predstavlja glavni izvor energije za rast i razvoj ploda. U posljednjem mjesecu graviditeta potrebe fetusa usljed intenzivnog rasta i razvoja rastu ato rezultira iscrpljivanjem rezervi majke i ketozom. Ona je osobito esta u mlije nih krava u peripartalnom razdoblju jer organizam nije sposoban istovremeno podr~avati razvoj ploda i stvoriti rezerve za nastupajuu laktaciju. Va~no je napomenuti kako proteini ne mogu prijei placentalnu barijeru, ve se razla~u na aminokiseline koje prolaze vezane na specifi ne nosa e. Iznimku predstavljaju imunoglobulini koji iako su proteini ipak mogu prei placentalnu barijeru kod placente endoteliochorialis (meso~deri) i placente hemochorialis (primati i glodavci). Svi ostali proteini se resintetiziraju u plodu od aminokiselina koje su stigle preko placente. Jednako tako ni lipidi ne prelaze placentalnu barijeru ve placenta hidrolizira trigliceride i fosfolipide majke koji razgraeni prelaze placentu i potom se ponovno sintetiziraju u lipide ploda. Veliki peptidni hormoni kao ato su tireoideostimulirajui hormon (TSH), adrenokortikostimulirajui hormon (ACTH), hormon rasta, inzulin i glukagon takoer ne mogu prei placentu ali zato steroidi, tiroidni hormoni i kateholamini (epinefrin i norepinefrin) bez poteakoa prelaze iz majke u plod. Vitamini i minerali takoer mogu prelaziti placentalnu barijeru ali s razli itim uspjehom. Vitamini topivi u vodi (B kompleks i C) lako prelaze, dok vitamini topivi u mastima (A,D, E i K) bitno te~e. Osim toga, hranjive tvari ulaze u plod i putem pinocitoze i fagocitoze. Horion tvori areole oko izlaza uterusnih ~lijezda i direktno apsorbira njihov sekret. Uz korisne, placenta mo~e u plod prenijeti i neke potencijalno atetne tvari. Tako primjerice etilni alkohol, ~iva, fosfor i mnoge druge toksi ne tvari lagano prolaze iz cirkulacije majke u plod. Jednako tako prelaze i razli iti opijati, barbiturati, amfetamin i razli iti antibiotici. Mnoge od tih tvari su izrazito teratogene i mogu oatetiti plod, izazvati malformacije u rastu i razvoju ploda ili prouzro iti abortus. Sli an u inak imaju i razli iti mikroorganizmi koji takoer mogu penetrirati u plod i izazvati infekciju. Osobito su esti virusi poput herpes virusa kao i specifi ne i nespecifi ne bakterije poput brucella, campilobactera i mnogih drugih kao i razli itih protozoa. Slika 3: Prikaz goveeg fetusa u placenti, gravidnost oko 90 dana; vidljivi kotiledoni, mjesta na kojima se placenta ve~e na karunkule majke tvorei placentome odgovorne za transport i interakcije cirkulacije ploda i majke  2.2.3. PLACENTA KAO ENDOKRINI ORGAN TIJEKOM GRAVIDNOSTI Placenta ima vrlo va~nu endokrinu ulogu u odr~avanju gravidnosti prakti ki od svog nastanka pa sve do poroda. Ugrubo mo~emo (prema Sengeru, 2003) endokrinu funkciju placente podijeliti na 5 razli itih poddjelova i to: stimulacija funkcije jajnika prvenstveno ~utog tijela odr~avanje gravidnosti utjecaj na fetalni rast i razvoj stimulacija funkcije mlije ne ~lijezde poticaj za porod Endokrina funkcija placente razlikuje se takoer od vrste do vrste pa tako kod kobile imamo tvorbu konjskog horionskog gonadotropina eCG (equine chorionic gonadotropin) a kod primata ljudskog horionskog gonadotropina hCG (human chorionic gonadotropin) koji predstavljaju ekstrahipofizarne gonadotropine. Zanimljivo je napomenuti da iako nisu fizioloaki prisutni u ostalih vrsta domaih sisavaca, a rije  je o peptidnim hormonima, svejedno mogu uzrokovati i dapa e koriste se za izazivanje folikulogeneze i ovulacije (krave, ovce, koze, kunike). U svakom slu aju eCG se lu i iz endometrijskih aaica a primarno djeluje kao luteotropin koji stimulira funkciju primarnog ~utog tijela kod kobile. Primarno ~uto tijelo je ono koje je nastalo nakon ovulacije koja je rezultirala koncepcijom. Akcesorna ~uta tijela koja su takoer va~na za odr~avanje gravidnosti u kobila nastaju od folikula koji su posljedica stimulacije upravo s eCG koji inducira nove ovulacije 40.-70. dana gravidnosti u kobile. ECG takoer inducira luteinizaciju antralnih folikula koji nisu ovulirali ali takoer prelaze u akcesorna ~uta tijela. Mogli bismo zaklju iti kako eCG ima vrlo zna ajan pozitivni utjecaj na sekreciju progesterona u kobila. Zanimljivo je napomenuti da aplikacija eCG poti e folikulogenezu u krava, ovaca i kuniki a da aplikacija eCG kobilama ne izaziva takav efekt i uope ne djeluje kao FSH. Kod primata se lu i hCG koji poput eCG potje e iz stanica trofoblasta horiona koji se po inje lu iti prakti ki im blastocista izgubi zonu pelucidu. Stoga ga koristimo u testovima za dokazivanje trudnoe jer je prisutan u krvi i mokrai trudnih ~ena ve 8-10 dana nakon koncepcije. Uloga hCG je podr~avanje funkcije ~utog tijela u prijelazu iz corpus luteum periodicum u corpus luteum graviditatis. Placenta takoer sintetizira i lu i steroidne hormone progesteron i estrogen. Progesteron je klju an za odr~avanje gravidnosti jer stimulira sekreciju uterusnih ~lijezda koje prehranjuju plod prije placentacije, a bitan je jer blokira kontrakcije miometrija i tako prevenira abortus. Estrogen je osobito va~an pred kraj gravidnosti jer poma~e prilikom pripreme za porod. Va~no je istaknuti kako izvor progesterona tijekom gravidnosti varira od vrste do vrste. Tako npr. kod ovce, kobile i ~ene placenta u potpunosti preuzima funkciju ~utog tijela pa luteoktomija kod ovce gravidne viae od 50 dana nee imati nikakvog u inka dok e kod krave gravidne manje od 8 mjeseci kao i kod krma e ili kunike tijekom itave gravidnosti dovesti do abortusa odnosno prijevremenog poroda. Va~no je napomenuti da i kod onih vrsta kod kojih placenta preuzme dominantnu funkciju u sintezi progesterona, svejedno na jajniku ostaje ~uto tijelo koje lu i progesteron prakti ki do poroda. Uz steroidne hormone placenta lu i joa itav niz polipeptida. Kod veine sisavaca placenta redovito lu i placentalni laktogen kojeg joa nazivamo i somatomamotropin. On je prisutan i mo~e se dokazati u serumu krave nakon 160. dana gravidnosti. Njegova funkcija je u pravilu dvojaka:on djeluje poput STH-somatotropnog hormona tako ato regulira i poti e rast fetusa te tako ato stimulira razvoj mlije ne ~lijezde (laktogeni efekt). Placentalni laktogen je dokazan kod atakora, miaeva, ovaca, krava i ljudi. Kod nekih vrsta prevladava prva a kod drugih druga navedena funkcija. Naprimjer kod ovaca i ljudi placentalni laktogen je mnogo zna ajniji kao stimulans za razvoj mlije ne ~lijezde nego za razvoj ploda dok je kod krava obrnuto. Smatra se da je razina sinteze placentalnog laktogena (PL) nasljedno svojstvo koje se prenosi preko mu~jaka pa se smatra da se kod mlije nih ovaca mo~e selekcijom ovnova odmah i direktno popraviti proizvodnja mlijeka u stadu. U svakom slu aju PL je produkt divovskih binuklearnih stanica koje lu e joa jedan vrlo zna ajan glikoprotein a to je PAG (pregnancy associated glicoprotein) kojeg mo~emo dokazati u serumu krave ve 24. dana nakon koncepcije. Njegova sinteza je karakteristi na za divlje i domae pre~iva e. Oba proteina i PAG i PL mogu se dokazati u cirkulaciji majke jer se oslobaaju iz sekretornih granula prilikom tvorbe stani nog sincicija. Placenta u ljudi, konja, pasa, ma aka, svinja i majmuna lu i joa i relaksin kojeg nema kod krava i njegova je funkcija kod krava dvojbena. Mogue je da ga krave uope niti ne sinteziraju. 2.3 SPECIFI NOSTI ENDOKRINOLOGIJE GRAVIDNOSTI U KRAVE Kod krave je tijekom itave gravidnosti ~uto tijelo na jajniku glavni izvor progesterona. Placenta takoer sintetizira manje koli ine progesterona. U svakom slu aju luteoktomija ili ovariotomija kod krave gravidne do 200 dana gravidnosti sigurno e dovesti do abortusa (Senger, 2003; Arthur, 1996). Tijekom posljednjeg mjeseca gravidnosti placenta i jajni ka stroma mogu kompenzirati funkciju ~utog tijela pa se graviditet nastavlja. Koncentracija progesterona je do 18 dana nakon ovulacije prakti ki identi na kod gravidne i negravidne krave. Nakon toga kod negravidne krave progesteron naglo pada i po inje novi ciklus dok kod gravidne takoer dolazi do laganog pada ali koncentracija ubrzo ponovno po inje rasti. Koncentracija progesterona ostaje visoka tijekom itave gravidnosti a po inje opadati 20-30 dana prije poroda. Koncentracija estrogena u opoj cirkulaciji je niska tijekom prve dvije treine gravidnosti i konstantno je ispod 100 pg/ml ali naglo naraste nakon 250. dana a vrhunac dosegne 5-7 dana pred porod - 7 ng/ml estron sulfata i 1,2 ng/ml estrona u krvi (Thatcher i sur.,1980). Do naglog pada estrogena dolazi 8h prije poroda pa sve do neposredno nakon poroda. Kod krave FSH i LH ostaju na bazalnoj razini itavu gravidnost i nema vidljivih fluktuacija u koncentraciji. Prolaktin ostaje nizak itavu gravidnost ali raste pred porod kada sa bazalne koncentracije od 50-60 ng/ml naraste na 320 ng/ml. Ponovno pada na bazalnu vrijednost vraa se 30h nakon poroda (Hoffman, 1973) Bovini placentalni laktogen (bpl) je prisutan u cirkulaciji nakon 160. dana a maksimalnu vrijednost od 1000 ng/ml dosegne oko 200. dana te ostaje visok do poroda (Bolander i sur. 1976). 2.4 LABORATORIJSKA, KLINI KA I ULTRAZVU NA DIJAGNOSTIKA RANE GRAVIDNOSTI U PRE}IVA A Metodologija koja se primjenjuje u dijagnostici gravidnosti uvelike se razvila u posljednjih 70 godina. Uz ve klasi ni znak izostanka estrusa u svih pre~iva a, rektalnu pretragu u krava i transabdominalnu palpaciju u malih pre~iva a danas se primjenjuju suvremene laboratorijske metode koje slu~e otkrivanju ~ivog embrija i maj inskog prepoznavanja gravidnosti, potom i ultrazvu na pretraga gravidnosti. Najbolja je naravno kombinacija svih pobrojanih metoda. Pri tome je va~no naglasiti kako nam je cilj ato prije otkriti negravidne jedinke, kako bismo ih im prije vratili u ciklus i ponovno osjemenili. Graviditet je u krava najranije mogue potvrditi odnosno isklju iti tri dana nakon umjetnog osjemenjivanja na osnovi postojanja EPF (early pregnancy factor) a sasvim pouzdane rezultate mo~emo dobiti ve 7-8 dana nakon osjemenjivanja (Adams i Jardon, 1999; Threlfall i Bilderbeck, 1999). Va~no je naglasiti da je EPF mogue utvrditi i iz krvi i iz mlijeka pa iako je metoda joa u stadiju intenzivnog razvoja, na tr~iatu su dostupni komercijalni kitovi pogodni za brzu i jednostavnu dijagnostiku. Velika mana navedene metode je ato iako dokazuje prisutnost embrija u jajovodu, odnosno maternici plotkinje, zapravo ne ukazuje na njegovu sposobnost pre~ivljavanja u maternici. Naime, kako navode Threlfall i Bilderbeck (1999) prisutnost embrija u organizmu majke ni im ne garantira da e se on zaista i razviti i opstati u maternici. Rana ultrazvu na dijagnostika gravidnosti u krava mogua je ve sa 9 dana (Pierson i Ginther, 1984), ali pouzdani dokaz da je plod ~iv imamo sa 24 dana kad su vidljivi otkucaji srca ploda (Goddard, 1994.). Od laboratorijskih metoda u prvom mjesecu gravidnosti u krava na raspolaganju nam stoje i razli iti trofoblast proteini (BPSP-bovine pregnancy specific proteins, PAG-pregnancy associated glycoproteins) koji se koriste u dijagnostici gravidnosti i rane embrionalne smrtnosti od sredine devedesetih (Zoli i sur., 1992, Beckers i sur., 1995, Szenci i sur., 1996, Karen i sur., 2002, Prvanovi i sur. , 2003) koje mo~emo mjeriti na osnovi RIA i ELISA metode. Njihova je prednost ato se mogu koristiti i retrogradno kao dokaz postojanja ~ivog vitalnog embrija kao i za razlikovanje embrionalne smrtnosti od izostanka koncepcije. Njihov je nedostatak ato zbog relativno duga kog bioloakog polu~ivota u opoj cirkulaciji ostaju prisutni u krvi joa 70 dana nakon poroda pa kod rane dijagnostike gravidnost nakon kratkog servis perioda je dosta teako razlikovati PAG prethodne od novonastale gravidnosti (Szenci, 2002). Na osnovi PAG profila mo~emo ve u prvom mjesecu gravidnosti utvrditi pojavu blizanaca (Dobson i sur., 1993, Patel i sur. 1995) kao i razli ite anomalije u razvoju (Ectors i sur., 1997, Szenci i sur., 1998). Od ostalih metoda rane laboratorijske dijagnostike koristi se odreivanje progesterona u krvi i mlijeku (Heap i sur. 1969, Robertson i Sardas, 1971). To nije dokaz postojanja embrija ve samo o uvane funkcije ~utog tijela. Sve navedene metode opisane u krava primjenjuju se i u malih pre~iva a. Kod ovaca se u ranoj dijagnostici koristi joa i rozeta inhibirajui test (RIT) koji slu~i za dokazivanje ranog faktora gravidnosti-EPF (early pregnancy factor) u prvom tjednu gravidnosti a prvi su ga opisali Morton i sur (1979). U ovce se gravidnost pomou navedene metode mo~e utvrditi 24h nakon parenja. Danas se intenzivno istra~uje mogunost primjene komercijalnih ELISA kitova za EPF u ovaca, dok se RIA kit za EPF pokazao nepouzdanim i neprakti nim (Sasser i Ruder, 1987). U ovaca se mo~e koristiti i vaginalna biopsija kao metoda rane dijagnstike, iako se ova metoda polako napuata (Richardson, 1972; Makek, 1976). Slika 4: Dijagnostika gravidnosti 72h nakon osjemenjivanja (RIT-rozeta inhibicijski test pomou ranog faktora gravidnosti (EPF-early pregnancy faktor)   Slika 5 a,b,c,d,: Ultrazvu ni nalaz gravidne krave nakon 3, 5, 7 i 9 tjedana gravidnosti a) tri tjednab) 5 tjedana EMBED Word.Picture.8 c) 7 tjedanad) 9 tjedana 2.5 ULOGA PAG U MAJ INSKOM PREPOZNAVANJU GRAVIDNOSTI I ODR}AVANJU GRAVIDNOSTI 2.5.1Blokiranje maj inog imunosnog odgovora tokom gravidnosti: implikacije za imunosupresiju Pre~ivljavanje sisavaca u maternici je direktno vezano za imunosupresiju koja omoguuje histoinkompatibilnom embriju da opstane u maternici. Presudnu ulogu u navedenoj imunosupresiji igra novootkriveni enzim indoleamin 2,3-dioxigenaza (IDO) koji se naziva joa i triptofan razgraujui enzim. On je zadu~en za rano maj insko prepoznavanje gravidnosti prije same placentacije i razvoja fetusa. Stanice koje posjeduju IDO kataboliziraju triptofan i spre avaju lokalnu aktivnost T limfocita i induciraju njihovu smrt. Tema ovog poglavlja biti e objaanjavanje uloge stanica koje lu e indoleamin 2,3-dioxigenazu (IDO) u regulaciji lokalnog imunosnog odgovora majke putem T limfocita tokom gravidnosti. Sama reakcija dobro je prou ena u miaeva, a smatra se da ima vrlo sli an tijek i u ostalih vrsta sisavaca. Lokalna imunosupresija posredstvom T limfocita nije do kraja razjaanjena ali se smatra da mo~e djelovati dvojako: ili tako da T limfociti uope ne prepoznaju stanice koje sadr~e antigen ili tako da doe do prepoznavanja stanica koje sadr~e antigen od strane T limfocita, ali da posljedi no ne doe do nikakve reakcije. U ovom drugom slu aju dolazi do aktivacije T limfocita ali i do uplitanja mehanizama koji spre avaju daljnju ekspanziju i diferencijaciju T limfocita. U te mehanizme su upleteni razli iti enzimi, makrofagi, dendriti ne i endotelijalne stanice i CD4+ stanice koje otpuataju CD25 koji suprimira periferni odgovor T stanica. U osnovi se joa uvijek ne zna ato odreuje hoe li aktivirani T limfociti djelovati tako da e uniatiti strane bjelan evine ili e jednostavno biti same blokirane i uniatene kao ato je to slu aj tokom rane gravidnosti. Fetus se u svakom slu aju od imunosne reakcije majke brani na tri razli ita na ina: fizi kim odvajanjem tkiva fetusa od tkiva majke, imunoloakom nezreloau samog fetusa i imunosupresijom majke (Dosogne i sur, 1999). U svakom slu aju je u ljudi i miaeva dokazano da T limfociti majke prepoznaju strano tkivo fetusa odnosno fetalne MHC (major histocompatibility complex) aloantigene tokom gravidnosti, a stanice senzibilizirane fetalnim proteinima mogu se dokazati u ~ena desetljeima nakon gravidnosti (Austin, 1999). Tim viae je interesantan mehanizam blokade ve senzibiliziranih i aktiviranih T limfocita. Pokusi invitro su dokazali da prisutnost veih koli ina esencijalne aminokiseline triptofana mo~e pozitivno utjecati na imunosni odgovor T-limfocita, dok je prisutnost stanica koje su lu ile IDO koji je katabolizirao triptofan dovela do blokade imunosnog odgovora T limfocita (Ahima i sur.,1998). Iz toga se dade zaklju iti da bi stanice koje lu e IDO mogle i u ~ivom organizmu blokirati imunosnu reakciju na lokalnoj razini tako ato limitiraju raspolo~ivu koli inu triptofana. Takoer se mo~e zaklju iti da je za aktivaciju T limfocita dovoljna niska razina triptofana. Pokusi na miaicama koje su nosile alogene fetuse a kojima su aplicirani inhibitori IDO izazvali su poba aje, dok u miaica koje su nosile singene fetuse nisu izazvali poba aje. Iz toga se dade zaklju iti kako IDO regulira lokalnu reakciju T limfocita ali nema nikakvog djelovanja na B limfocite. Time se ujedno objaanjava kako i zaato mo~emo pronai antitijela majke za tkiva fetusa a da svejedno ne dolazi do odbacivanja tijekom gravidnosti. U svakom slu aju stanice koje sadr~e IDO dokazane su na povraini embrija i endometrija ve neposredno nakon implantacije. Istra~ivanja su nadalje pokazala da se 50-100 takvih stanica redovito nalazi na pola puta izmeu materni nih ~lijezda i ektoplacentalnog konusa u miaica gravidnih 7,5 dana. Pokazalo se da se ne radi niti o makrofagima niti o NK (natural killer cells) majke. Tri dana kasnije njihov je broj jako porastao i okru~ili su itav embrij. Ukoliko je gravidnim miaicama apliciran inhibitor IDO, aktivirao se sustav komplementa, koji je uz pomo osna~enih i diferenciranih T limfocita, eliminirao embrij. Zanimljivo je i primjetiti da embrij treba triptofan kao esencijalnu aminokiselinu za svoj rast i razvoj, a IDO ga blokira i razgrauje. Embrij to najvjerojatnije prevladava tako da relativno brzo transportira triptofan u stanice a zadaa je stanica koje sadr~avaju IDO i okru~uju embrij da fizi ki onemogue pristup T-limfocitima do dovoljnih koli ina triptofana i da paralelno razgrade sav raspolo~ivi triptofan koji nije preuzeo embrij a mogao bi poslu~iti T limfocitima. Pokusi na ljudskim i miajim T limfocitima invitro su pokazali da nedovoljna koli ina triptofana u mediju izaziva prestanak proliferacije T-limfocita i dovodi do apoptoze. Pokusi sa dodavanjem i uklanjanjem ostalih esencijalnih aminokiselina u mediju su pokazali da oni nemaju nikakvu zna ajniju ulogu u proliferaciji T-limfocita koji su osjetljivi jedino na koncentraciju triptofana osobito u prijelazu iz G1 u S fazu. Zanimljivo je dodati da ve navedene stanice koje sadr~e IDO takoer slu~e i kao nosioci antigena za izazivanje tvorbe protutijela i ako blokiramo IDO stanice ne samo da e doi do odbacivanja ploda, ve e doi i do izostanka humoralne imunosti na antigene ploda. Stanice koje sadr~e IDO djeluju i kao regulatori upalne reakcije takoer preko katabolizma triptofana. Njihov izostanak je zabilje~en kod pacijenata oboljelih od AIDS-a i HTLV, a izaziva otpuatanje neurotoksina, demenciju i mijelopatije. Iako je sigurno da su IDO stanice prisutne kod upale, nije u potpunosti jasno da li IDO stanice poma~u ili atete kod upalne reakcije. S jedne strane IDO stanice mogu biti korisne utoliko ato ubla~uju i reguliraju snagu imunosnog odgovora organizma, usporavaju rast tumora i proliferaciju T limfocita, ali s druge strane mogu biti i atetne jer katabolizam triptofana rezultira tvorbom metabolita koji mogu biti toksi ni po zdrave stanice. IDO je heme-sadr~avajui enzim koji katabolizira prvi ograni avajui korak u oksidativnoj degradaciji triptofana. Za nasljeivanje IDO u sisavaca dobro je sa uvan samo jedan gen u koji je odgovoran i za tvorbu upalnih medijatora kao interferon . Naime, linijske kulture stanica podrijetlom od ljudskih mijeloidnih, stromalnih i ostalih tumora otpuataju IDO kad dou u kontakt sa interferonom , dok je u svje~im kulturama tih istih stanica nemogue dokazati IDO. LPS i upalni citokini IL1 i TNF djeluju sinergisti ki sa interferonom  kako bi doveli do daljnjeg porasta IDO u ljudskim dendriti nim i Hella stanicama, dok antiupalni citokini IL4 IL10 i TGF inhibiraju IDO. U glodavaca je pojava IDO limitirana samo na pojedina tkiva kao ato su limfni vorovi, djelovi crijeva, slezena, timus, plua, epididimis i fetoplacentalni spoj. Pokazalo se takoer da je pojava IDO u miaeva vezana na upalnu reakciju jednako kao i na porast progesterona u serumu ali to joa nije do kraja dokazano. Ono ato je sigurno to je da IDO stanice putem katabolizma triptofana moduliraju lokalni imuni odgovor organizma a sve uzro no-posljedi ne veze i odnose treba joa istra~iti. Odreenu ulogu u blokiranju T-limfocita majke sasvim sigurno imaju i PAG (pregnancy associated glycoproteins). Oni su biokemijski srodni tripsinu i ostalim proteazama ali se pokazalo da nemaju sposobnost proteoliti ke aktivnosti (Guruprasad i sur., 1996). Prema istra~ivanju kojeg su proveli Xie i sur. (1991) i Roberts i sur. (1996) postoji pretpostavka da PAG imaju primarno endokrinu funkciju. Nadalje, pokazalo se da imaju sposobnost vezivanja na specifi ne stani ne receptore majke nakon ato su otpuateni iz stanica trofoblasta. PAG takoer imaju sposobnost vezivanja i transporta razli itih proteina ovisno o svojoj strukturi i upravo se na osnovi vezivanja razli itih proteina mo~e razlikovati razli ite PAG molekule. Roberts i sur (1996) su dokazali da PAG ima sposobnost kompeticije sa MHC za peptide prezentirane kao antigen i na taj na in interferiraju s aktivacijom T-limfocita. Najvjerojatnije je da PAG djeluju lokalno na povrini placente jer se neki od njih javljaju predominantno na mononuklearnim trofoblast stanicama koje nisu invazivnog karaktera, a PAG se mogu dokazati i na placenti kobila i svinja kod kojih nema erozije uterusnog epitela a pristup u cirkulaciju majke je ograni en. Endokrinu ulogu PSBP (pregnancy associated binding proteins) koji spada u PAG, objasnio je Austin i sur (1999) koji su otkrili da porast BPSP izaziva otpuatanje  kemokina koji je identi an sa granulocitnim kemotakti kim proteinom-2 (GCP-2). Identi an efekt izaziva i interferon- tijekom prvih nekoliko tjedana koncepcije, a takoer djeluje lokalno. Nadalje,  kemokini mogu regulirati adheziju, upalnu reakciju i angiogenezu tijekom rane implantacije a mogu biti upleteni i u restituciju endometrija nakon poroda (Perenyi, 2002). Jednako tako Del Vecchio i sur.(1990, 1995) su dokazali da PAG stimulira lutealnu funkciju jajnika i porast P4 u serumu. Isti autori su na pokusima invitro dokazali da PAG 1 djeluje na koncetraciju PGF2, PGE2 i oksitocina i time indirektno utje e na blokadu luteolize. PAG djeluje na koatanu sr~ i tako utje e na sintezu polimorfonuklearnih neutrofila ato izaziva promjene u imunosnom odgovoru organizma (fagocitoza, oksidacijski procesi) ato su Dosogne i sur. (1999) i dokazali. Va~no je naglasiti i to da su istra~ivanja molekularne biologije PAG dokazala da se razli ite podvrste PAG javljaju tijekom razli itih faza gravidnosti. Tako se boPAG 1,6 i 7 javljaju tijekom srednjeg i kasnog graviditeta, dok se boPAG 2,3,4 i 5 javljaju na samom po etku, ve oko 25. dana gravidnosti. (Garbayo i sur. 2000, Green i sur. 2000). Kod vrsta koje imaju placentu epiteliochorialis kao ato su kobila i krma a PAG javlja se samo eqPAG-1, poPAG-1, poPAG-2 i to na povraini horiona (Green i sur., 1999). 2.5.2 Uloga placente i binuklearnih stanica u razvoju i odr~avanju rane gravidnosti u goveda Placenta je specijalizirano, ekstraembrionalno tkivo koja igra izuzetno va~nu ulogu u razvoju i odr~avanju gravidnosti. Rije  je multifunkcionalnom organu koji ima respiratornu, nutritivnu, epurativnu, endokrinu i imunoloaku funkciju. Placenta goveda je sinepitelihorijalna odnosno sindezmohorijalna a horionske resice su lokalizirane u ograni enim podru jima okruglog do ovalnog oblika koje nazivamo kotiledoni, dok itavu placentu nazivamo placenta kotiledonaria. Kotiledoni sa placente stapaju se sa proliferiranim slojem endometrija-karunkulima i zajedno ine placentome. Broj kotiledona u krava varira od 70-150 smjeatenih u 5 redova kako u gravidnom tako i u negravidnom rogu (3 reda po sredini rogova i po dva reda na prednjem dijelu rogova). Kotiledoni goveda su konveksni i peteljkasti. Interkarunkularno podru je endometrija (paraplacenta) ostaje ~ljezdano aktivno tijekom gravidnosti dok je karunkularno podru je tijekom gravidnosti bez ~ljezdane aktivnosti (neglandularno). Specifi nost placente goveda je populacija binuklearnih stanica (BNC). One nastaju iz mononuklearnih stanica trochoderma procesom nuklearne mitoze bez citoplazmatske mitoze. Najmlae binuklearne stanice nalaze se duboko u trohodermu a tijekom zrenja migriraju prema povraini maternice (Wooding i sur. ,1983) Binuklearne stanice se pojavljuju neposredno prije implantacije (Wango i sur.,1990) a u trohodermu goveda mogu se prepoznati od 16-17 dana nakon koncepcije. Takve stanice predstavljaju otprilike 15-20% svih stanica trohoderma od implantacije sve do nekoliko dana pred porod kada njihov broj rapidno opada (Wooding 1983, Wooding i sur., 1986). Morfometrijske kvantitativne studije su pokazale da 20% binuklearnih stanica migrira preko horionskih resica u endometrij gdje se stapaju sa stanicama endometrija i tvore maj inske divovske trinuklearne stanice. (Wooding 1984, Wooding and Beckers 1987). Takve trinuklearne stanice su kratkog ~ivotnog vijeka jer nakon otpuatanja svojih sekretornih granula posredstvom egzocitoze bivaju resorbirane od trohoderma (Wooding and Wathes, 1980) Mlade BNC sadr~avaju mali volumen guste citoplazme ispunjene ribosomima koja nakon citoplazmatske reorganizacije u okruglu ili ovalnu stanicu nema kontakta s bazalnom membranom ili atipi nim trohodermalnim spojem. U procesu zrenja razvija se endoplazmatski retikulum i veliko Golgijevo tijelo. BNC su uklju ene u tvorbu fetomaternalnog sincicija koji je esencijalan za uspjeanu implantaciju i posljedi no rast placentoma. Upravo ova populacija stanica va~na je za sintezu i sekreciju razli itih proteina i steroidnih hormona. Binuklearne stanice su izvor placentalnih laktogena (Verstegen i sur., 1985, Wooding i sur., 1992) Istra~ivanja koja su uklju ivala imunohistokemiju dokazala su da da se proteinski hormoni u BNC sintetiziraju kroz ednoplazmatski retikulum a tijekom zrenja spremaju u granule (Wooding, 1981; Wooding i Beckers, 1987). Pokazalo se da su binuklearne stanice odgovorne za sintezu PAG, BPSP (Reimers i sur.,1985; Zoli i sur., 1992; Atkinson i sur, 1993). Binuklearne stanice takoer su upletene i u sintezu i sekreciju progesterona (Wango i sur., 1991; 1992; Wooding i sur., 1996). Izolirane binuklearne stanice su sposobne sintetizirati prostaglandine (Reimers i sur.,1985) va~ne u parakrinoj regulaciji sinteze progesterona. Slika 8: Prikaz fuzije endometrijskih binuklearnih stanica i stanica trofoblasta u divovoske trinuklearne stanice (prema Woodingu i Wathesu, 1980) 1-stanica trofoblasta; 2-binuklearna stanica u endometriju; 3- trinuklearna stanica; 4-otputanje PAG egzocitozom u krvotok majke; 5-trinuklearna stanica nakon egzociteze u endometriju, 6- trinuklearna stanica u korionu  Slika 9: Prikaz migracije stanica trofoblasta u endometrij majke i otputanje PAG u krvotok majke (Senger, 2003)  2.5.3. Odreivanje trofoblast proteina (PAG, PSPB I PS60) pomou RIA metode Preparati PAG, PSPB i PSP60 koriateni su za pripremu antiseruma na kuniu ato je omoguilo razvoj specifi ne RIA reakcije za detekciju navedenih spojeva u bioloakim tekuinama (Sasser i sur. (1986), Humblot i sur., 1988), Mialon i sur., 1993 i Zoli i sur., 1992) Specifi na dvostruka-antitjelo (double-antibody) reakcija RIA omoguila je mjerenje bPAG u ekstraktima placente, fetalnom serumu, plodnim vodama, te u serumu i plazmi gravidnih krava. Navedenu metodu modificirao je Zoli sa suradnicima 1992. godine. Minimalna mjerljiva koncentracija iznosila je 0,2 ng/ml seruma. Koncentracija PAG bila je zna ajno viaa u maj inom nego u fetalnom serumu, kada je mjerena istovremeno ato potvruje da se ovi proteini lu e prvenstveno u krvotok majke. To je sasvim suprotno od placentalnog laktogena koji je predominantno visok u cirkulaciji fetusa nakon 160. dana gravidnosti, a gotovo je nemjerljiv i jedva prisutan u cirkulaciji majke. Isti autor navodi kako se u 20% juvenilnih junica i negravidnih krava takoer mo~e pronai mjerljiva razina PAG iako je ona daleko ispod fizioloakih granica. PAG je mogue otkriti u serumu majke otprilike u vrijeme kada se trofoblast definitivno ve~e za stijenku maternice (Zoli i sur.,1992; Sasser i sur. 1986). Tako je otprilike 22. dana nakon oplodnje razina PAG u prosjeku iznosila 0,380,13 ng/ml seruma (Perenyi, 2002) Potom dolazi do polaganog porasta PAG tijekom rane i srednje gravidnosti. Sinclair i sur. (1995) su opisali krivulju eksponencijalnog rasta bPAG izmeu 23. i 85. dana gravidnosti u krava nakon UO. Dramati an porast uslijedio je tokom posljednjih 10 dana gravidnosti, kada je koncentracija PAG dostigla 2462,41017,9 ng/ml. Mialon i sur. Koji su pratili razinuPSP60 (pregnancy specific proteins 60) otkrili su da je ukupna koncentracija svih proteina specifi nih za gravidnost narasla 80-200x tijekom posljednja dva tjedna gravidnosti. Sli an trend primijetili su i Sasser i sur. (1986) koji su pratili razinu PSPB tokom itave gravidnosti a utvrdili dramati an porast u predpartalnom razdoblju. Poslije poroda redovito dolazi do pada koncentracije PAG sve do jedva mjerljive razine 10020 dana poslije poroda (Zoli i sur., 1992). Bioloaki polu~ivot PAG i ostalih glikoproteina ploda iznosi 7,4 do 9 dana (Kiracofe i sur., 1993; Mialon i sur., 1993; Ali i sur.,1997). Smatra se da bieksponencijalna jednad~ba najbolje opisuje nestanak PAG iz periferne cirkulacije (Ali i sur., 1996). Manje razlike uo ene su izmeu pojedinih pasmina ato se objaanjava razli itom distribucijom odnosno metabolizmom PAG izmeu plazme i jetre ovisno o tome pripada li krava tovnom ili mesnom tipu odnosno da li je metabolizma optereen proizvodnjom velikih koli ina mlijeka ili nije. Tako je primjerice razina PAG zna ajno viaa kod krava i junica pasmine Hereford nego kod krava i junica pasmine Holstein (Zoli i sur, 1992). Osim toga, pokazalo se da razina PAG je zna ajno viaa kod krava pasmine Charolais nego u krava pasmine Holstein i normandijskih pasmina krava tokom posljednja dva mjeseca gravidnosti (Mialon i sur.,1993) Pokazalo se da koncentracija PAG raste ukoliko je napravljen embriotransfer, osobito ako se radi o transferiranju embrija druge pasmine ili ak vrste (Beckers i sur., 1998; 1999). Takoer je primjeeno kako na razinu PAG zna ajno (p<0,01) utje e stadij gravidnosti i broj plodova. Naime, krave koje su nosile blizance imale su bitno viae razine PAG i PSPB tokom itave gravidnosti uz izuzetak posljednjih 10 dana gravidnosti (Patel i sutr.,1995, 1997,1998, Dobson i sur., 1993). Osobito je zna ajna razlika u razini PAG uo ena nakon i oko 40. dana gravidnosti izmeu krava koje su nosile jedan plod od onih koje su nosile blizance. (Vasques i sur. 1995) Smatra se da je tako velika razlika kod krava koje nose blizance posljedica dva mjesta na kojima se odvija placentacija (Patel i sur. 1997). Postoji takoer pozitivna korelacija izmeu koncentracije PAG u posljednja dva tjedna pred porod i poroajne te~ine teleta (Patel i sur. 1995) 2.5.4. Primjena trofoblast proteina (PAG) u dijagnostici gravidnosti u krava Iako je sigurno da PAG mo~e poslu~iti u ranoj dijagnostici gravidnosti u krava, razli iti autori navode razli ite vrijednosti i pozdanost metode. Tako Mialon i sur. (1993) tvrde da je dijagnostika gravidnosti na osnovi razine PAG u serumu 100% pouzdana nakon 27. dana gravidnosti, dok Humblot i sur.(1988) i Delahaut i sur. (1996) tvrde da je metoda 100% pozdana tek nakon 30 dana. U svakom slu aju mo~emo rei da je metoda nakon 24. dana pouzdana preko 90%, a osobito je korisna za negativnu dijagnostiku, odnosno njome je mogue pouzdano isklju iti graviditet. Sasser i sur. (1986) su opisali primjenu PAG nas komercijalnom stadu od 102 krave tovne pasmine pri emu su ustvrdili da je dijagnostika gravidnosti spomou PAG bila pouzdanija i primjenjivija ranije od rektalne pretrage. Primjena RIA testa za PAG omoguuje pouzdano isklju ivanje negravidnih krava izmeu 26. i 30. dana gravidnosti s to noau od 90% (Humblot i sur., 1988) Isti autori dobili su 100% to ne rezultate u svih krava koje su bile gravidne viae od 30 dana. Pozitivna dijagnostika gravidnosti bila je pouzdana u 74% krava i 90% junica gravidnih 28 dana Va~no je naglasiti da na to nost nalaza tijekom rane gravidnosti mo~e utjecati i PAG iz prethodne gravidnosti odnosno duljina servis perioda. Ukoliko je interval izmeu poroda i uzimanja krvi iznosio viae od 115 dana, tada je pouzdanost metode 28. dana gravidnosti iznosila 84% za pozitivnu dijagnozu, a preko 90% za negativnu dijagnozu (Mialon i sur.,1994). Prilikom terenskog istra~ivanja Zoli i sur. (1992) su mjerili razinu bPAG u serumu junica nakon embriotransfera na 287 Holstein Frizijskih junica. Ustanovili su da je metoda dala pouzdano pozitivne rezultate u 93,4% slu ajeva a a pouzdano negativne rezultate u 97,9% slu ajeva. Ukupna pozdanost testa iznosila je 94,6%. Krv je uzimana 35 dana nakon estrusa, a nalaz je potvren rektalnom pretragom sprovedenom 45. dana. Nadalje, koncentracija bPAG u serumu mo~e se uspjeano koristiti u odreivanju starosti fetusa u sparenih tovnih junica. Naime, to je est problem ukoliko se radi o haremskom dr~anju kod kojeg nemamo pouzdane podatke o estrusu niti o gravidnosti, a ~elimo znati kada e doi do teljenja (Sinclair i sur. 1995). U istra~ivanju koje su proveli Skinner i sur (1996) na 233 junice pri emu su komparirali razli ite metode dijagnostike gravidnosti. Utvrdili su da je PAG metoda izbora jer pokazuje osjetljivost od 100%, specifi nost 93%, da je pouzdanost pozitivne dijagnoze 97% i pouzdanost negativne dijagnoze 100% te ukupnu pouzdanost 98%. Sli ne rezultate dobio je i Szenci (1998) koji je usporedio to nost dijagnostike gravidnosti posredstvom ultrazvuka i PAG RIA testa izmeu 27. i 31. dana gravidnosti. Nakon usporedbe dvije laboratorijske metode RIA bPAG i RIA bPSPB pouzdanost oba testa bila je sli na i iznosila oko 95% (Szenci i sur., 1995) pa ih u daljnjem razmatranju neemo izdvojeno promatrati. Isti autor je primjetio da na osnovi rezultata teljenja u skupini krava koju je pregledavao (n=243) nije bilo zna ajne razlike izmeu ultrazvu ne pretrage, bPAG i bPSPB. Glavnu poteakou prilikom la~no pozitivnog nalaza kod primjene PAG u ranoj dijagnostici gravidnosti predstavlja uzimanje uzoraka unutar 100 dana nakon poroda. Naime, u tom su periodu glikoproteini ploda zaostali od prethodne gravidnosti joa uvijek prisutni u cirkulaciji majke, upravo zbog svog relativno duga kog bioloakog polu~ivota. Stoga je primjena PAG u ranoj dijagnostici gravidnosti ograni ena u terenskim uvjetima (Perenyi, 2002) ali je izuzetno korisna jer omoguuje otkrivanje rane embrionalne smrtnosti. 2.5.5. Praenje gravidnosti i dijagnostika rane embrionalne smrtnosti na osnovi PAG profila Osim za dijagnostiku gravidnosti PAG se mo~e koristiti i kao marker gravidnosti, jer ukoliko doe do uginua ploda dolazi do postepenog pada PAG u serumu majke. Stoga je vrlo korisno viaekratno uzimati krv gravidnim ~ivotinjama kako bismo otkrili razliku izmeu embrionalne smrtnosti i izostanka koncepcije. Ipak i tu postoje ograni enja jer je nemogue otkriti uginue embrija ukoliko on nije doaao u kontakt sa sluznicom endometrija i placentacija nije otpo ela (Mialon i sur., 1993). Kao ato je poznato PAG molekule su produkt stanica trofoblasta i smatra se da mogu poslu~iti kao pokazatelj vitalnosti embrija, te pomoi u otkrivanju i dokazivanju ranih anomalija placente, embrionalne smrtnosti i abortusa (Ectors i sur. 1996; Drame i sur., 2000). Junice koje su izgubile plod 16-35 dana nakon UO imale su razli ite krivulje PAG i sve su ukazivale na gubitak ploda. Ovo se objaanjava razli itim vremenom uginua embrija (Mialon i sur., 1993). Takoer se pokazalo ako do uginua ploda doe oko 50. dana tada razina PAG u serumu joa neko vrijeme raste jer su stanice trofoblasta uklju ene u placentaciju pa su nakon uginua ploda starog 40-50 dana joa neko vrijeme aktivne i ~ive, ato su otkrili Mialon i sur.(1993) koji su paralelno ultrazvu no pratili gravidnost od 17. dana pa do 50. dana i to tako da jsu svakih 72h pregledavali krave i uzimali uzorke krvi i potom pratili daljnji graviditet. Isti su autori utvrdili da kod manjeg postotka gravidnih krava takoer mo~e doi do prolaznog pada razine PAG oko 40. dana no uskoro dolazi do ponovnog porasta koji je konstantan sve do poroda. Humblot i sur.(1988) prou avali su u inkovitost BPSP testa i odreivanja progesterona u mlijeku 24. dana nakon UO i ustvrdili da u 60% slu ajeva dolazi do povratka u ciklus jer do koncepcije nije niti doalo ili je pak embrij uginuo prije antiluteoliti kog signala, dakle kada je bio mlai od dva tjedna. Isti autori su utvrdili da ne postoji korelacija izmeu razine BPSP i razine progesterona tokom itave gravidnosti. Takoer su ustvrdili da postoji zna ajna razlika izmeu razine BPSP izmeu skupine gravidnih krava i skupine krava koje su pretrpjele embrionalnu smrtnost od 24. do 35. dana nakon UO. U svom radu oni nisu mogli predvidjeti pojavu embrionalne smrtnosti na osnovi prethodnih nalaza ali su zaklju ili da mjerenje progesterona paralelno s PSPB u serumu krava, mo~e pomoi u otkrivanju kako razli iti imbenici mjerljivi u serumu kronoloaki mogu utjecati na pojavu rane embrionalne smrtnosti kao i njene posljedice. Tokom nekoliko godina Ectors i sur.(1996; 1997) su istra~ivali razinu PAG nakon invitro oplodnje i kloniranja. Oni su uzimali krv svakih tjedan dana i otkrili da je mogue na osnovi PAG profila odrediti kada je doalo do embrionalne odnosno fetalne smrtnosti. Isti autor je u drugom istra~ivanju (1996b) ustvrdio kako je na PAG profil utjecalo i samo podrijetlo embrija pa su junice koje su nosile embrije dobijene invitro imale viau razinu PAG u kasnijoj gravidnosti od junica koje su nosile embrij nakon nuklearnog transfera. To se objaanjava veim postotkom morfoloakih anomalija, poput placentalne hipertrofije, hidroamniona, i hidatiforme molare, te porasta broja i promjera kotiledona koji se javljaju nakon nuklearnog transfera. Autori smatraju da navedene anomalije placente nastaju kao posljedica nedovoljnog zrenja oocita u vrijeme oplodnje, te manjeg promjera folikula koji nisu rezultirali kompetentnim oocitama ili su rezultirali samo djelomi no kompetentnim oocitama, forsiranim u razvoju. (Ectors i sur., 1997) Jednako tako Heyman i sur. (2002) otkrili su da primateljice koje imaju hidroalantois imaju izrazito poviaene vrijednosti BPSP koji prelazi 400ng/ml seruma. Isti autori utvrdili su sna~nu pozitivnu korelaciju izmeu anomalija u razvoju placente i nepravilnog PAG profila. Ectors i sur. (1997) koji su pratili PAG profile kod junica koje su pobacile u ranoj gravidnosti (tokom 7 i 8 tjedna gravidnosti) te u kasnoj fazi (nakon 158 dana gravidnosti) potvruju ranije navedene tvrdnje. Patel i sur. (1997) su opisali da krave koje su iznijele plod do termina, ali je plod bio nakazan i uginuo (konkretno, radilo se o schistosomi reflexus) u pravilu imaju nepravilan porast PAG pred sam porod, koji ne odgovara fizioloakim vrijednostima. Jednako tako ~iv ali slabo vitalan plod izneaen do termina, kada je uginuo pred porod ili u porodu, izaziva neuobi ajen porast PAG u drugoj treini gravidnosti. Openito mo~emo rei da se u veterinarskoj praksi podjednako mo~emo koristiti mjerenjem PAG i BPSP u bioloakim tekuinama u potvrdi i praenju gravidnosti (Szenci, i sur., 1998) Naime, krave kod kojih je nakon prvog ultrazvu nog pregleda postavljena dijagnoza da su gravidne a nakon drugog ultrazvu nog nalaza da su negravidne, u pravilu imaju i pad PAG u opoj cirkulaciji izmeu navedena dva pregleda. Za razliku od njih, potvreno gravidne krave na osnovi oba ultrazvu na pregleda imaju vrlo visoke razine PAG i progesterona u opoj cirkulaciji. Ukoliko je doalo do embrionalne smrtnosti tada dolazi do pada PAG, a razina progesterona ostaje visoka u 75% slu ajeva (Perenyi, 2002). Osim toga, Szenci i sur.(2000) koji su prou avali PAG profil tokom uginua ploda u raznim fazama gravidnosti, ustanovili su da je mogue utvrditi vrijeme uginua ploda ve na osnovi PAG profila. Pri tome su naglasili kako je zbog vrlo velikih individualnih varijacija u razini PAG u serumu gravidnih krava i junica izuzetno bitno ustanoviti zna ajan pad PAG u serumu kako bismo sa sigurnoau mogli tvrditi da se radi o embrionalnoj smrtnosti. 2.6 BIOKEMIJSKA SRODNOST LEPTINA I OSTALIH CITOKINA VEZANIH ZA GRAVIDNOST (PREGNANCY ASSOCIATED CYTOKINES) Root: scop Class: All alpha proteins Fold: 4-helical cytokines Family: citokini dugog lanca Interferoni/interleukin-10 (IL-10) Protein: Leptin interferon-tau Placentalni laktogen Slika 9: Shematski prikaz molekule leptina (a) i interferona  (b) a) leptinb) interferon -  HYPERLINK "http://www.3dchem.com/molecules.asp?ID=154" \l "#"  INCLUDEPICTURE "http://www.3dchem.com/imagesofmolecules/Leptin.jpg" \* MERGEFORMATINET  2.7. ULOGA LEPTINA U REPRODUKCIJI Od otkria leptina u 1995. godini jako se puno istra~uje njegova funkcija u domaih i divljih sisavaca. Do sada se otkrilo jako puno o ulozi leptina u murida i primata, dok se o ulozi leptina u drugih vrsta sisavaca puno manje zna. Osobito je zanimljiva i nedovoljno istra~ena uloga koju leptin ima u domaih pre~iva a a vezana je za pozitivan i negativan energetski balans , utjecaj ciklusa, gravidnosti i laktacije. Leptin je 167-amino kiselinski peptid molekularne mase od 16 kD kojeg sintetizira i lu i masno tkivo (Cunningham i sur.,1999; Capro i sur. 1999). Za leptin se ispo etka smatralo da utje e na energetsku ravnote~u tako ato djeluje centralno da bi inhibirao unos hrane (Zhang i sur.,1994) te djeluje na imunosni sustav i neuro-endokrinu osovinu. Receptori za leptin pronaeni su u mozgu, masnom tkivu, hipofizi, jajnicima i placenti (Lyn i sur.,2000; te Dyer i sur.,1997). Dokazano je da leptin djeluje na unos glukoze i sintezu glikogena preko inzulina, a koncentracija leptina u serumu direktno modulira neuro-hormonalne interakcije (Ngyen van Ly,2003). Pokusi na miaevima su pokazali da ukoliko im apliciramo leptin dolazi do poveane potroanje kisika, poja ane lokomotorne aktivnosti i poviaene tjelesne temperature. Aplikacija leptina izaziva porast koncentracije LH, FSH i testosterona u plazmi, a kod intra-cerebro-ventrikularne aplikacije (ICV) stimulira sekreciju LH i estrogena u primarno ovariektomiziranih atakorica (Yu i sur.,1997). Leptin igra zna ajnu ulogu u svim fazama reprodukcije. Pokusi na miaevima i ljudima dokazali su da je upravo porast koncentracije leptina signal koji inicira po etak spolnog dozrijevanja i uspostavu ciklusa (Ahima i sur.,1997). 2.7.1. Uloga leptina u odr~anju reproduktivne sposobnosti Aplikacija leptina u ~enke miaeva izaziva porast LH i FSH u serumu, dok kod muakih miaeva izaziva porast mase testisa i broja spermija u ejakulatu (Barash i sur, 1996). Nadalje, opetovana parenteralna aplikacija leptina rezultira ovulacijom, gravidnoau i kasnije porodom nakon osjemenjivanja u ~enki miaeva (Chehab i sur., 1996) Istra~ivanja na rezus majmunicama su pokazala da konstantna parenteralna aplikacija leptina izaziva porast FSH i LH u plazmi ( Finn i sur., 1998). Isti autor je dokazao da aplikacija leptina stimulira neurone koji otpuataju GnRH u hipotalamusu a ujedno direktno utje e na hipofizu tako ato je poti e na sintezu FSH i LH. Leptin svakako djeluje i na spolni ciklus u pre~iva a. Zanimljivo je napomenuti da je aplikacija rekombinantnog ov jeg leptina izazvala porast LH u serumu neovisno o dozi kod podhranjenih intaktnih krava (Amstalden i sur., 2002), dok je u sli nom pokusu izvedenom na podhranjenim ovarijektomiziranim kravama reakcija ovisila o dozi. Tako je aplikacija leptina u koncentraciji 0,2-2 ng/ml izazvala stimulaciju sekrecije LH, dok vee ili manje koncentracije nisu imale nikakvog efekta. (Zieba i sur.2003). Za razliku od krava intracerebroventrikularne infuzije rekombinantnog ov jeg leptina u podhranjenih i normalno hranjenih ovaca i ailje~ica ne utje u na razine FSH i LH u serumu (Henry i sur., 1999, Morrison i sur.,2001). `toviae, Morrison i sur. (2002) su otkrili da niti intravenska aplikacija leptina nikako ne utje e na porast LH u serumu u mladih ailje~ica koje se nalaze u fazi intenzivnog rasta i dozrijevanja. Stoga bi se moglo rei da specifi an utjecaj leptina na sekreciju gonadotropnih hormona u pre~iva a ovisi o dobi jedinke (prije i poslije dostizanja spolne dozrelosti) kao i o jajnicima (velika razlika izmeu ovariektomiziranih i intaktnih ~ivotinja). 2.7.2. Utjecaj leptina na po etak puberteta Sa sigurnoau je dokazano da leptin regulira po etak puberteta u miaeva (Ahima i sur. 1997, Chehab i sur. 1997), atakora (Gruaz i sur., 1998) te u ljudi (Matkovi i sur, 1997). U ljudi je dokazano da razina leptina intenzivno raste tijekom puberteta u oba spola, a potom kod ~ena ostaje visoka dok kod muakaraca opada jer porast razine testosterona djeluje kao inhibitor sekrecije leptina (Horlick i sur.,2000). Time je ujedno mogue objasniti razlike u koncentraciji leptina u jedinki iste grae i mase a razli itog spola (Havel i sur., 1996, Rosenbaum i sur.,1996). Parenteralna aplikacija leptina ubrzava pojavu i tijek puberteta u miaeva (Ahima i sur.,1997, Chehab i sur.,1997) i atakora (Gruaz i sur. 1998). Koncentracija leptina u mlije nih krava linearno je rasla nakon 16. tjedna prije prve ovulacije u jednogodianjih junica (Garcia i sur.,2002) a bila je u direktnoj linearnoj korelaciji sa tjelesnom masom (Diaz-Torga i sur., 2001). Za razliku od njih, Block i sur. (2003) nisu utvrdili porast leptina prije prve ovulacije, ato objaanjavaju izostankom povoljnog utjecaja metaboli kih faktora u njihovih junica odnosno tvrde da leptin djeluje stimulativno na pojavu prve ovulacije, ali sam po sebi ne mo~e izazvati prvu ovulaciju. 2.7.3.Uloga leptina u gravidnosti Istra~ivanja sprovedena na gravidnim ~enama utvrdila su da leptin po inje rasti tijekom rane gravidnosti, nastavlja rasti tokom druge i tree treine i ostaje visok sve do poroda (Mukherjea i sur., 1999). Sli no je i kod atakora (Amico i sur.,1998; Garcia i sur. 2000), potom miaeva (Tomimatsu i sur., 1997) te ovaca (Erhardt i sur. 2001,; Forhead i sur. 2002). Svi autori su sla~u da je do porasta koncentracije leptina doalo zbog gomilanja masnog tkiva koje nastaje u trudnoi i paralelno poveanja mRNA ekspresije za leptin u masnom tkivu. Porast leptina u gravidnosti je zapravo paradoksalan jer je gravidnost stanje kada su poveane potrebe organizma za energijom i hranjivim tvarima pa bi bilo logi no da leptin pada. Va~no je napomenuti kako leptin tokom gravidnosti uglavnom mo~emo dokazati u neaktivnom vezanom obliku u opoj cirkulaciji ato vjerojatno mjenja njegovu bioraspolo~ivost odnosno poveava rezistenciju na leptin (Mounzih i sur., 1998, Seeber i sur. 2002). U krava koncentracija leptina u ranoj gravidnosti nije do kraja istra~ena dok su istra~ivanja koja su uklju ivala mjerenje razine leptina u srednjoj i kasnoj gravidnosti dokazala da dolazi do njegovog porasta (Block i sur.,2001). To je takoer neobi no imajui u vidu da je leptin hormon podrijetlom iz adipocita, koji se prete~no sintetizira u masnom tkivu te lu i u opu cirkulaciju u omjeru koji je upravo proporcionalan koli ini masnog tkiva. U fizioloakim koncentracijama (2-10 ng/ml serum) on poti e steroidogenezu jajnika i razvoj embrija ali ako je vrlo visoka (oko 100 ng/ml seruma) tada inhibira oboje. Koncentracija leptina zna ajno raste u gravidnih ovaca tokom rane gravidnosti (u prva dva tjedna) i smatra se da regulira invazibilnost citotrofoblasta tijekom implantacije i placentacije. Leptin djeluje na embrij tako ato poveava unos i metabolizam glukoze putem regulacije metabolizma i sinteze inzulina. Sve navedeno je dokazano u ljudi, miaeva, atakora i ovaca. Pokusi na ovcama su pokazali da porast koncentracije leptina u gravidnih ovaca potje e ispo etka od citotrofoblasta, potom od placente te kona no od masnog tkiva ploda. Koncentracija leptina u krvi ostaje poviaena tijekom cijele gravidnosti i rapidno opada nakon poroda, u ljudi, ovaca, miaeva i atakora. Leptin je po svojoj strukturi biokemijski vrlo srodan placentalnom laktogenu i interferonu-tau, ato ukazuje na mogunost da bi mogli imati i sli nu funkciju. Leptin tijekom gravidnosti u ljudi dokazano direktno utje e na: rast i razvoj zametka, feto-placentalnu angiogenezu, embrionalnu hepatopoezu te biosintezu hormona unutar feto-placentalnog spoja ploda i majke (Henson i Castracane,2000). Leptin spada u porodicu citokina-1, koja takoer uklju uje hormon rasta i leukemija inhibitorni faktor (LIF) (Tartaglia i sur.,1995). Pokazalo se da su i leptin i LIF bitni za mehanizam implantacije i angiogenezu (Vogiagis i Salamonsen,1999). Istra~ivanja su pokazala da se leptin nalazi vezan na vilozne endotelne stanice (Henson i sur, 2000) te da je koncentracija leptina zna ajno viaa u umbilikalnih vena nego u umbilikalnih arterija (Yura i sur.,1998), ato zajedno ukazuje na injenicu da je placentalni leptin va~an za fetalni rast i razvoj, premda se ne mo~e isklju iti niti uloga fetalnog leptina. U svinja je koncentracija leptina via u placentama koje opskrbljuju normalne fetuse od onih koje opskrbljuju retardirane fetuse, a koji su se nalazili u istoj maternici (Ashworth i sur., 1998). Placenta dokazano proizvodi leptin u mjerljivim koncentracijama u serumu takorica (Masuzaki i sur., 1997), babuna (Henson i Castracane, 2000) i imia (Zhao i sur.,2003). Za razliku od njih, u miica nema mjerljive razine placentalnog leptina (Gavrilova i sur., 1997; Tomimatsu i sur. 1997; Kronfeld-Schor i sur., 2001.; Zhao i sur., 2003) to je u svojim istra~ivanjima pobio jedini Hoggard i sur. (1997). Takoer i placenta krava, za razliku od placente ovaca lu i zanemarive koncentracije placentalnog leptina tokom itave gravidnosti (Bispham i sur., 2003) ato navodi na zaklju ak da je sinteza leptina u placenti strogo vrsno specifi na i razlikuje se ak i izmeu vrlo srodnih vrsta kao ato su primjerice pre~iva i. Takoer je va~no naglasiti kako je tvorba leptina u placenti regulirana neovisno o masnom tkivu, ato su u humanoj ginekologiji dokazali Bie i sur. (1997), pa je vrlo vjerojatno da je tvorba placentalnog leptina neovisna i u ostalih vrsta sisavaca tokom gravidnosti 2.7.4.Utjecaj leptina na funkciju jajnika Razli iti su autori u svojim istra~ivanjima neovisno doali do konstatacije kako leptin djeluje izravno na jajnike u svih vrsta domaih sisavaca. Tako primjerice u goveda granuloza stanice (Spicer i Francisco, 1997) i stanice thece (Spicer i Francisco, 1998) imaju receptore za leptin i razina steroidogeneze ovisi o leptinu. Kod svinja se pokazalo i da je direktno ovisna o koncentraciji leptina u serumu (Ruiz-Cortes i sur, 2003) istra~ivanja na atakorima pokazala su da parenteralna aplikacija visokih doza leptina (30 (g svakih 3h tijekom 15 h) negativno utje e na stopu ovulacije ali nema nikakav utjecaj na FSH niti LH (Duggal i sur, 2000). Istra~ivanja provedena na ~enama pokazala su da se leptin u ~ena lu i u mjese nim intervalima (Hardie i sur., 1997) i odreene fizioloake promjene u koncentraciji leptina su neophodne da bi se odr~ao ciklus na jajnicima (Laughlin i sur.,1997). Sli ne promjene u razini leptina vezano uz pojavu spolnog ciklusa zabilje~ene su u svinja (Lin i sur., 2000, Ruiz-Cortez i sur., 2000) i atakorica (Zamorano i sur., 1997) 2.7.5.Povezanost razine leptina i laktacije Kao ato je ve navedeno u ranijem poglavlju, razina leptina je poviaena tijekom gravidnosti i naglo pada nakon poroda. Visoka razina leptina povezana je s pripremama organizma za laktaciju i poveanim rezervama masnog tkiva ato rezultira i poviaenjem koncentracije leptina ato je dokazano u atakorica i krava (Woodside i sur., 2000, Block i sur., 2001). Nakon poroda i tokom laktacije dolazi do pada koncentracije leptina ato se objaanjava poja anim energetskim potrebama organizma tijekom laktacije i mobilizacijom masnih rezervi organizma. Pokazalo se da na razinu leptina tokom laktacije ni na koji na in ne utje e sisanje teleta niti intenzitet mu~nje (Brogan i sur., 1999), Pickavance i sur.,(1998).su primjetili da u krava pad leptina u serumu zbog negativnog energetskog balansa izaziva poja ani apetit ato rezultira obnavljanjem tjelesnih rezervi neophodnih za odr~avanje laktacije. Takoer je va~no naglasiti da leptin najvjerojatnije direktno utje e na razinu mamogeneze u krava i ovaca. Naime, dokazano je da u obje vrste postoje receptori u mlije noj ~ljiezdi na koje se tokom gravidnosti ve~e velika koli ina leptina, kao i da je intenzitet razvoja mlije ne ~lijezde tokom rane i srednje gravidnosti pod direktnim utjecajem razine leptina u serumu (Smith i Sheffield, 2002.; Bonnet i sur. 2002; Laud i sur., 1999) 3. MATERIJAL I METODE 3.1.Odabir plotkinja za istra~ivanje Krave koje su koriatene za ovo istra~ivanje bile su simentalske pasmine, u dobi 3-7 godina, redovito su koriatene u reprodukciji i pri tome nisu imale nikakvih poteakoa sa koncepcijom ni sa poroajem. Dr~ane su na 5 obiteljskih gospodarstava u Meimurskoj ~upaniji, u uzgojima koji su brojili od 8 do 20 krava. Prilikom provoenja istra~ivanja nije doalo do zna ajnijih promjena laktacijske krivulje (mjerljive u dnevnoj koli ini mlijeka) ni tjelesne kondicije (mjerljive putem BCS-body condition score-a). Sve su dr~ane na vezu, muzene strojno i hranjene mjeaavinom zelene i kukuruzne sila~e, koncentrata i mineralnim dodacima. Telile su se redovito svake godine, poroaji su im bili lagani i samostalni ili uz laganu pomo, a sve krave su bez ikakvih problema ponovno zabreale 70-90 dana nakon poroda. }ivotinje su kasnije, ovisno o rezultatima koncepcije odnosno ishodu gravidnosti premjeatene u jednu od 3 skupine. Krave koje su normalno iznijele plod do termina, a plod je na svim ultrazvu nim pregledima pokazivao znakove vitalnosti i viabilnosti predstavljale su pokusnu skupinu 1. Krave kojima je gravidost potvrena ultrazvu nom pretragom nakon 17.-24. dana od U.O. smatrane su gravidnom skupinom sve dok se na osnovi rektalne i ultrazvu ne pretrage nije ustvrdilo da je doalo do uginua ploda. Te su ~ivotinje kasnije praene zasebno, kao pokusna skupina 2-skupina embrionalne odnosno fetalne smrtnosti a na osnovi laboratorijskih pokazatelja (progesteron, PAG, leptin) sam pokuaala ustvrditi kada je doalo do smanjenog vitaliteta i posljedi no uginua ploda. }ivotinje koje su uzete u istra~ivanje a nisu koncipirale u navedenom roku poslu~ile su kao kontrolna skupina jer su bile istog pasminskog sastava i dobi, a dr~ane su u istim uvjetima kao krave iz pokusne skupine. Takve ~ivotinje su se iz razli itih razloga (visoka laktacija, subestrus, lutealne ciste) tiho gonile ili nisu niti uale u ciklus nakon poroda, pa nisu niti osjemenjene. Svejedno smo im oduzimali krv po istom re~imu kao i kravama iz ostalih skupina kako bismo dobivene parametre na kraju mogli usporediti i isklju iti sve ostale utjecaje na rezultate pretraga, osim graviditeta. Jednako tako smo i kasnije, ukoliko bismo tijekom provoenja istra~ivanja ( tijekom rektalne i ultrazvu ne kontrole gravidnosti) u bilo kojem trenutku ustvrdili da je doalo do poba aja ili uginua ploda, plotkinju prebacili u skupinu krava koje su pobacile odnosno u skupinu krava kod koje je doalo do rane embrionalne, embrionalne ili fetalne smrtnosti, ovisno o tome kada je doalo do uginua zametka. Istra~ivanjem smo sveukupno obuhvatili 74 krave, podijeljene u gravidnu (pokusna skupina 1), gravidnu koja je pobacila (pokusna skupina 2) i negravidnu, neosjemenjenu skupinu (kontrolna skupina). Sve krave koje su odabrane po gornjem kriteriju praene su od poroda pri emu su sve imale normalan puerperij i involuciju maternice. Prvi manifestni estrus je evidentiran i ultrazvu no potvren do 60 dana nakon poroda u veine krava. Potom su ~ivotinje ultrazvu no praene i osjemenjene u drugom manifestnom estrusu. Nakon toga su sve osjemenjene krave tretirane jednako. Ukoliko su ultrazvu ni pregledi 17. i 35. dana pokazali da je krava gravidna, ona je svrstana u skupinu gravidnih krava i praena do idueg teljenja. Ukoliko krava nije koncipirala od prvog osjemenjenja,a ponovno se tjerala, osjemenili smo je ponovno. Ukoliko ni tada nije doalo do koncepcije, krava je izba ena iz istra~ivanja, odnosno nije dalje osjemenjena ve je praena zajedno sa skupinom negravidnih, neosjemenjenih kontrolnih ~ivotinja. Svi uzorci krvi krava sumnjivih na pojavu embrionalne smrtnosti promatrani su kao zasebna skupina, kako bi se utvrdilo da li je zaista u kravi postojao ~ivi, vitalni plod. 3.2. Uzimanje krvi i ultrazvu no praenje gravidnosti nakon umjetnog osjemenjivanja Svim kravama nakon UO krv je redovito vaena ili iz kokcigealne vene ili iz vene jugularis pomou vacutainer sustava i to u jednakim razmacima. Pri tome je krv uzeta u SST epruvete i odmah po uzorkovanju izcentrifugirana na 3000 okretaja .Nakon toga je serum preliven u eppendorf kivete i smrznut na  20C do kona nih zajedni kih pretraga. Serumi su odle~ali razli ito vrijeme pohranjeni na navedenoj temperaturi, ali ukupno vremensko razdoblje od prvog uzorkovanja do posljednje pretrage iznosilo je manje od 6 mjeseci, pa nije utjecalo na kona ne rezultate. Svim smo kravama prilikom svakog uzimanja krvi uzeli ukupno po 4 ml seruma pohranjena u duplikatu u kivetama od 2 ml. Jedna kiveta poslu~ila je za odreivanje PAG (pregnancy associated glycoprotein) i progesterona, a druga za odreivanje koncentracije leptina u serumu. Krv je uzimana na sam dan osjemenjivanja (nulti dan) i potom svaka 72h sve do 40. dana nakon U.O. Prilikom vaenja krvi, kravama je fiksirana glava i vrat, a uz ~ivotinju je redovito bila vlasnica ili neka druga poznata osoba koja je umirivala kravu i pomagala kod obuzdavanja i sputavanja. Mirnim kravama je krv vaena iz repne vene ( v. coccigea) jer je to dokazano manje stresno za krave od vaenja iz v. jugularis, ali je neizvedivo kod nemirnih i plahih krava zbog opasnosti od ozljeivanja osoba koje sudjeluju u vaenju krvi. Krave su redovito klini ki pregledavane i praene prakti ki od poroda. 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Grafikon 8:Usporedba razine PAG1 za sve skupine krava  EMBED STATISTICAGraph  Analiza varijance takoer je pokazala da postoji signifikantna razlika (P>0,01) za srednje vrijednosti PAG 2 za sve tri skupine odnosno izmeu gravidnih krava i krava s embrionalnom smrtnoau, gravidnih krava i negravidnih krava te krava s embrionalnom smrtnoau i negravidnih krava. Grafikon 9:Usporedba razine PAG2 za sve skupine krava  EMBED STATISTICAGraph  Analiza varijance takoer je pokazala da postoji signifikantna razlika (P>0,05) za srednje vrijednosti PAG 3 za sve tri skupine odnosno izmeu gravidnih krava i krava s embrionalnom smrtnoau, gravidnih krava i negravidnih krava te krava s embrionalnom smrtnoau i negravidnih krava. Grafikon 10:Usporedba razine PAG3 za sve skupine krava  EMBED STATISTICAGraph  Analiza varijance za PAG 4 je pokazala da izmeu gravidnih krava(1) i krava s embrionalnom smrtnoau (2) postoji postoji visokosignifikantna razlika (p>0,01) jednako kao i i izmeu gravidnih i negravidnih krava (3) dok izmeu skupine krava s embrionalnom smrtnoau i negravidnih krava nema signifikantne razlike. Grafikon 11:Usporedba razine PAG4 za sve skupine krava  EMBED STATISTICAGraph  Analiza varijance za PAG 5 je pokazala da izmeu gravidnih krava(1) i krava s embrionalnom smrtnoau (2) postoji postoji visokosignifikantna razlika (p>0,01) jednako kao i i izmeu gravidnih i negravidnih krava (3) dok izmeu skupine krava s embrionalnom smrtnoau i negravidnih krava nema signifikantne razlike. Grafikon 12:Usporedba razine PAG5 za sve skupine krava  EMBED STATISTICAGraph  Analiza varijance za progesteron 1. je pokazala da izmeu gravidnih krava(1) i krava s embrionalnom smrtnoau (2) postoji signifikantna razlika (p>0,05) jednako kao i i izmeu gravidnih i negravidnih krava (3) dok izmeu skupine krava s embrionalnom smrtnoau i negravidnih krava nema signifikantne razlike. Grafikon 13:Usporedba razine progesterona 1 za sve skupine krava  EMBED STATISTICAGraph  Analiza varijance za progesteron 2. je pokazala da izmeu gravidnih krava(1) i krava s embrionalnom smrtnoau (2) postoji signifikantna razlika (p>0,05) jednako kao i i izmeu gravidnih i negravidnih krava (3) dok izmeu skupine krava s embrionalnom smrtnoau i negravidnih krava nema signifikantne razlike. Grafikon 14:Usporedba razine progesterona 2 za sve skupine krava  EMBED STATISTICAGraph  Analiza varijance za progesteron 3. je pokazala da izmeu gravidnih krava(1) i krava s embrionalnom smrtnoau (2) postoji signifikantna razlika (p>0,05) jednako kao i i izmeu gravidnih i negravidnih krava (3) dok izmeu skupine krava s embrionalnom smrtnoau i negravidnih krava nema signifikantne razlike. Grafikon 15:Usporedba razine progesterona 3 za sve skupine krava  EMBED STATISTICAGraph  Faktori linearne korelacije istra~ivanih parametara Faktori linearne korelacije istra~ivanih parametara promatrani su i komparirani po pojedinim skupinama krava (gravidne, negravidne i embrionalna smrtnost). Prikazani su pojedina no za svaku skupinu u tablicama koje slijede. Analiza linearnih korelacija u svim skupinama pokazala je da nema statisti ki zna ajnih korelacija izmeu pojedinih vrijednosti unutar skupine kao ni izmeu razli itih skupina. Faktori linearne korelacije istra~ivanih parametara Tablica 13: Korelacije u skupini gravidnih krava-signifikantne korelacije odatampane su crvenim brojkama: LEPT1LEPT2LEPT3LEPT4PAG1PAG2PAG3PAG4PAG5PROG1PROG2PROG3LEPT110,3313190,3745210,4236850,0784670,1296070,2042440,0146290,14656-0,20888-0,30823-0,15284LEPT20,33131910,863990,607621-0,003810,0596-0,02107-0,11086-0,00933-0,32778-0,30135-0,18894LEPT30,3745210,8639910,8697530,0237530,1935050,077957-0,03640,024278-0,2792-0,25437-0,10803LEPT40,4236850,6076210,86975310,2175380,2829860,1570340,0830150,118244-0,1662-0,19435-0,03512PAG10,078467-0,003810,0237530,21753810,594220,2952610,3103810,2558470,1639190,1659890,154765PAG20,1296070,05960,1935050,2829860,5942210,650480,4431230,3225380,1780240,0831540,079534PAG30,204244-0,021070,0779570,1570340,2952610,6504810,8212020,7312160,3058460,199810,17628PAG40,014629-0,11086-0,03640,0830150,3103810,4431230,82120210,8577570,3393970,2866030,246305PAG50,14656-0,009330,0242780,1182440,2558470,3225380,7312160,85775710,3495630,3217850,289687PROG1-0,20888-0,32778-0,2792-0,16620,1639190,1780240,3058460,3393970,34956310,9122570,815721PROG2-0,30823-0,30135-0,25437-0,194350,1659890,0831540,199810,2866030,3217850,91225710,928545PROG3-0,15284-0,18894-0,10803-0,035120,1547650,0795340,176280,2463050,2896870,8157210,9285451 Tablica 14: Korelacije u skupini krava s embrionalnom smrtnoau-signifikantne korelacije odatampane su crvenim brojkama: LEPT1LEPT2LEPT3LEPT4PAG1PAG2PAG3PAG4PAG5PROG1PROG2PROG3LEPT110,6549130,6304890,6087340,1603540,01414-0,05434-0,08056-0,065640,3746510,1027780,222729LEPT20,65491310,9340280,698718-0,084210,117739-0,08625-0,11637-0,115650,069492-0,02404-0,11685LEPT30,6304890,93402810,798634-0,059440,10659-0,05637-0,10526-0,097850,2379340,010153-0,05216LEPT40,6087340,6987180,79863410,092350,2108130,0707020,0719030,0950540,239206-0,122460,123122PAG10,160354-0,08421-0,059440,092351-0,22951-0,28146-0,2809-0,2593-0,03943-0,317260,034189PAG20,014140,1177390,106590,210813-0,2295110,7372940,6818340,628542-0,11128-0,12273-0,06546PAG3-0,05434-0,08625-0,056370,070702-0,281460,73729410,8978010,859384-0,15354-0,12087-0,148PAG4-0,08056-0,11637-0,105260,071903-0,28090,6818340,89780110,993184-0,31474-0,26064-0,2321PAG5-0,06564-0,11565-0,097850,095054-0,25930,6285420,8593840,9931841-0,30879-0,26214-0,24159PROG10,3746510,0694920,2379340,239206-0,03943-0,11128-0,15354-0,31474-0,3087910,4952360,758331PROG20,102778-0,024040,010153-0,12246-0,31726-0,12273-0,12087-0,26064-0,262140,49523610,131959PROG30,222729-0,11685-0,052160,1231220,034189-0,06546-0,148-0,2321-0,241590,7583310,1319591 Tablica 15: Korelacije u skupini negravidnih krava-signifikantne korelacije odatampane su crvenim brojkama: LEPT1LEPT2LEPT3LEPT4PAG1PAG2PAG3PAG4PAG5PROG1PROG2PROG3LEPT110,8602180,7617530,598647-0,235880,377488-0,24446----0,229322-0,429780,051747LEPT20,86021810,9225090,691160,0970630,052167-0,19935-----0,03998-0,51873-0,16948LEPT30,7617530,92250910,8952750,103309-0,02507-0,10353-----0,00987-0,47026-0,10611LEPT40,5986470,691160,89527510,003157-0,077650,149483----0,166411-0,350870,119357PAG1-0,235880,0970630,1033090,0031571-0,08323-0,08323-----0,35909-0,14509-0,37419PAG20,3774880,052167-0,02507-0,07765-0,083231-0,05882----0,071712-0,10045-0,11359PAG3-0,24446-0,19935-0,103530,149483-0,08323-0,058821----0,071712-0,09310,271887PAG4--------------1--------PAG5----------------1------PROG10,229322-0,03998-0,009870,166411-0,359090,0717120,071712----10,2092710,681462PROG2-0,42978-0,51873-0,47026-0,35087-0,14509-0,10045-0,0931----0,20927110,186924PROG30,051747-0,16948-0,106110,119357-0,37419-0,113590,271887----0,6814620,1869241 5. RASPRAVA Za uspjeano praenje koncepcije i ranog tijeka gravidnosti neophodno je procijeniti sve faktore koji mogu eventualno utjecati i dovesti do izostanka koncepcije ili pak do rane embrionalne smrtnosti. Jednako tako va~no je uo iti kakvo je stanje u cjelokupnom uzgoju kao i u kolikom postotku se javila rana embrionalna smrtnost a u kolikom se radi o izostanku koncepcije. Samo na taj na in mo~emo sa sigurnoau tvrditi kada je i kako doalo do uginua embrija. Poznato je da je rano pre~ivljavanje embrija u jednakoj mjeri ovisno i o karakteristikama ploda i karakteristikama majke. Upravo je rana embrionalna smrtnost jedan od mehanizama kojima organizam majke pravovremeno odbacuje avitalne ili pak geneti ki deformirane embrije koji nisu sposobni izazvati maj inski odgovor na gravidnost. U tu smo svrhu mi u naaem istra~ivanju uzeli u obzir sve potencijalno va~ne imbenike kao i njihove meusobne odnose kako bismo ne samo pokuaali otkriti ranu embrionalnu smrtnost i razlu iti je od izostanka koncepcije, ve i pokuaali objasniti kako je do nje doalo. Jednako tako, bio nam je cilj otkriti uzgoje u kojima se javlja rana embrionalna smrtnost te otkriti zajedni ke egzogene i endogene imbenike koji su ju prouzro ili. Sve navedeno je od izuzetne va~nosti kako bismo razlu ili primarno neplodne krave (one koje su nesposobne koncipirati ) od onih kod kojih je doalo do uginua embrija. Svrha ovog rada bila je istra~iti kakva je uloga trofoblast proteina i leptina u fiziologiji rane gravidnosti u goveda kao i kolika je mogunost njihove primjene u dijagnostici embrionalne smrtnosti. Naime, od presudnog je zna aja u ranoj dijagnostici gravidnosti u goveda, pravovremeno izdvojiti negravidne plotkinje te ih im prije vratiti u ciklus i ponovno osjemeniti, kako bi potencijalni gubici u proizvodnji bili im manji. Osobito je va~no razlikovati primarno neplodne jedinke od onih kod kojih je doalo do uginua embrija, kako bi se otkrili uzroci navedenih pojava a ~ivotinje nakon odgovarajueg tretmana ili lije enja im prije ponovno zabreale. Kako bismo mogli pratiti ranu gravidnost od ranije poznatih i uvrije~enih metoda koristili smo redovite rektalne preglede, transrektalnu ultrasonografiju i odreivanje progesterona u serumu. Kako bismo mogli kontrolirati dobivene rezultate, sve krave u pokusu smo pratili do poroda i na osnovi podataka o telenjima, mogli smo definitivno potvrditi odnosno isklju iti gravidnost u promatranih grla.Ultrazvu na pretraga se ve 17. dana pokazala pouzdana u 95% slu ajeva ato se sla~e s rezultatima koje je objavio Szenci (1995). Potvrda odnosno isklju enje gravidnosti nakon ultrazvu ne pretrage sprovedene 35. dana nakon UO dala je pouzdanost od 100%, jednako kao ato je opisao i Goddard (1994). Statisti ka analiza rezultata dobivenih za razinu progesterona 12., 21. i 35. dana nakon UO pokazala je da postoji signifikantna razlika (p>0,05) izmeu gravidnih krava i krava s embrionalnom smrtnoau jednako kao i i izmeu gravidnih i negravidnih krava. Izmeu skupina krava s embrionalnom smrtnoau i negravidnih krava nema signifikantne razlike. Mo~emo zaklju iti da je na osnovi razine progesterona u serumu mogue razlu iti gravidne od negravidnih krava ali nije mogue dijagnosticirati embrionalnu smrtnost. Kako bismo mogli pratiti razvoj embrija i uo iti pojavu embrionalne smrtnosti, pristupili smo praenju razine PAG 24., 30., 34., 40 i 45. dana nakon umjetnog osjemenjivanja. Nakon statisti ke obrade pokazalo se da rezultati variraju ovisno o vremenu uzorkovanja. Analiza varijance je pokazala da postoji signifikantna razlika (P>0,01) za srednje vrijednosti PAG 24., 30 i 34. dana nakon UO za sve tri skupine odnosno izmeu gravidnih krava i krava s embrionalnom smrtnoau, gravidnih krava i negravidnih krava te krava s embrionalnom smrtnoau i negravidnih krava. Suprotno tome,analiza varijance za razinu PAG 40. i 45.-tog dana nakon UO je pokazala da izmeu gravidnih krava i krava s embrionalnom smrtnoau postoji visokosignifikantna razlika (p>0,01) jednako kao i izmeu gravidnih i negravidnih krava dok izmeu skupine krava s embrionalnom smrtnoau i negravidnih krava nema signifikantne razlike. Iz toga se mo~e zaklju iti da je embrionalna smrtnost vidljiva na osnovi drasti nog pada razine PAG u serumu majke 7,5-9 dana nakon uginua embrija, koliko iznosi i bioloaki polu~ivot PAG u prvom mjesecu gravidnosti, kao ato su opisali i Kiracofe i sur. (1993), Mialon i sur. (1993) te Ali i sur. (1997). Takoer mo~emo zaklju iti i da je u naaem istra~ivanju ako promatramo skupinu krava s embrionalnom smrtnosti, 98% krava izgubilo embrij kad su bile gravidne 17-24 dana ato predstavlja kriti no razdoblje za pre~ivljavanje embrija. Navedenu tvrdnju mo~emo potkrijepiti i nalazom progesterona i komparacijom razine progesterona izmeu skupina iz koje je vidljivo da postoji signifikantna razlika izmeu gravidnih krava i ostale dvije skupine, dok izmeu negravdnih krava i krava kod kojih je doalo do embrionalne smrtnosti nema signifikantne razlike u koncentraciji progesterona za niti jedno mjerenje. To se apsolutno sla~e s podacima iz literature pa tako Mialon i sur. (1993) navode kako do embrionalne smrtnosti vidljive na osnovi PAG profila naj eae dolazi u krava gravidnih 16-24 dana, dok Humblot i sur. (1988) navode kako do uginua embrija u pravilu dolazi do 24. dana. S njima se sla~u i rezultati istra~ivanja koje su objavili Ectors i sur. (1996.) te Drame i sur. (2000) koji su prou avali mogunosti dijagnostike pre~ivljavanja ploda u maternici krave na osnovi PAG profila i obojica su zaklju ili kako je do drasti nog pada u razini PAG u krava kod kojih je plod uginuo doalo 24. do 35. dan od koncepcije, ato je ako broju dana kada su utvrdili pad PAG oduzmemo 7,5 do 9 dana koliko iznosi bioloaki polu~ivot T2 za PAG, identi no rezultatima koje smo mi dobili u naaem istra~ivanju. Do postepenog padanja koncentracije PAG proteina dolazi stoga ato su u serumu gravidne krave prisutne razli ite molekule PAG, razli itih funkcija i bioloakog polu~ivota, pa se, kako navodi Green i sur. (2000) nakon 27. dana javljaju u serumu razli ite podvrste PAG koje mogu dati la~no visok, pozitivan nalaz. Mogue je i da stanice trofoblasta opstanu i egzistiraju u endometriju a samim tim i lu e PAG proteine joa neko vrijeme nakon uginua embrija, kao ato je opisao Mialon i sur. (1993). Svi navedeni autori meutim isti u da neposredno po uginuu embrija, krivulja PAG naglo pada ili dobiva nepravilan oblik ato omoguuje to no odreivanje vremena uginua embrija. Analiza varijance u naaem istra~ivanju je pokazala da nema signifikantne razlike izmeu skupina za srednje vrijednosti leptina niti za jedno uzorkovanje izmeu svih skupina. Na osnovi naaih rezultata vidljivo je da je doalo do porasta koncentracije leptina u serumu nekih od gravidnih krava u naaem istra~ivanju, ali na~alost ne postoje statisti ki zna ajne korelacije izmeu gravidnosti i porasta razine leptina u serumu. Jednako tako je va~no naglasiti da je kod velike veine ~ivotinja u sve tri skupine zapravo doalo do laganog pada leptina u serumu ato je posljedica uvoda u laktaciju i porasta lu enja dnevne koli ine mlijeka po kravi. Pad razine leptina tijekom uvoda u laktaciju ve je opisan i dokazan u istra~ivanjima koja su objavili Woodside i sur., (2000) te Block i sur.,(2001). Za razliku od nas koji smo istra~ivanje izveli na kravama pasmine simentalac, oni su istra~ivali na kravama pasmine holstein i pritom su dobili sli an rezultat kao i mi. Naime i u njihovim i u naaim istra~ivanjima doalo je do pada koncentracije leptina tokom uvoda u laktaciju ato mo~emo objasniti poja anim energetskim potrebama organizma tijekom laktacije i mobilizacijom masnih rezervi organizma. Takoer je va~no naglasiti da leptin najvjerojatnije direktno utje e na razinu mamogeneze u krava i ovaca. Naime, dokazano je da u obje vrste postoje receptori u mlije noj ~lijezdi na koje se tokom gravidnosti ve~e velika koli ina leptina, kao i da je intenzitet razvoja mlije ne ~lijezde tokom rane i srednje gravidnosti pod direktnim utjecajem razine leptina u serumu (Smith i Sheffield, 2002., Bonnet i sur. 2002, Laud i sur., 1999) ato bi se trebalo odraziti i na porast koncentracije leptina u krvi tokom rane i srednje gravidnosti. No problem u objektivnom procjenjivanju razine leptina u serumu predstavlja injenica ato se navedena dva suprotna efekta javljaju istovremeno. Rana gravidnost izaziva porast leptina,a laktacija istovremeno izaziva njegov pad. Upravo to superponiranje uvoda u laktaciju i rane gravidnosti onemoguuje objektivnu procjenu profila leptina u krvi. Naime, iako je viae nego vjerojatno da je leptin direktno upleten u pre~ivljavanje embrija u krava i da njegova koncentracija direktno ovisi o vitalnosti embrija, to je nemogue dokazati samo direktnim mjerenjem leptina u serumu. Za sada nema na ina da se razlu i utjecaj laktacije na koncentraciju leptina od utjecaja gravidnosti na istu jer se odvijaju istovremeno. Jednako tako mo~emo zaklju iti kako pri tome razina laktacije ne igra zna ajnu ulogu, jer je rezultat isti neovisno o tome da li se radi o visokomlije nim kravama ili o kravama kombinirane pasmine kao ato je simentalac koje smo mi koristili u naaem istra~ivanju. `toviae, pokazalo se da iako su krave simentalske pasmine grublje i robustnije konstitucije i s manjim promjenama kondicije tijekom laktacije od visokomlije nih holstein krava (ato onemoguuje direktno preslikavanje primjene odreivanja laktacijske krivulje BCS na simentalke) ipak se razina laktacije zna ajno odrazila na koncentraciju leptina u serumu ato zna i da je mobilizacija tjelesnih rezervi masti vea nego ato je to vidljivo na osnovi odreivanja BCS (body condition scorea). Nemogue je direktno povezati utjecaj rane gravidnosti goveda na razinu leptina u serumu, kao ato je to slu aj kod ostalih pre~iva a, iako sve ukazuje na to da on postoji. Zato bi bilo preporu ljivo iskuaati pratiti razinu leptina u serumu krava uz odreivanje BCS s paralelnim ultrazvu nim mjerenjem sloja potko~ne masti te istovremenim praenjem razine leptina u gravidnih i negravidnih krava kako bismo otkrili eventualnu povezanost i posljedi nu primjenu mjerenja leptina u serumu kao pokazatelja odr~ivosti rane gravidnosti u krava. Jednako tako valja naglasiti, obzirom da je u naae istra~ivanje uklju en relativno veliki broj ~ivotinja (n=74) podjednake proizvodnje, pasminskog tipa i na ina dr~anja, da su vrijednosti leptina ukoje smo izmerili u serumima krava bile vrlo raznolike za pojedina mjerenja i nisu ovisile niti samo o laktaciji niti samo o gravidnosti, smatramo da postoji vrlo zna ajan individualni faktor koji utje e na razinu leptina u krava pa na~alost ne mo~emo koristiti leptin kao direktni indikator rasta i razvoja ploda u krava u prvih mjesec dana nakon koncepcije. To naravno ne zna i da leptin ne igra ulogu u pre~ivljavanju embrija kod krava sli nu onoj kod ostalih pre~iva a (ovca, koza, deva, ljama,koauta) ve samo zna i da je razina leptina u serumu ovisna o itavom nizu drugih imbenika koji mogu i ne moraju nu~no biti povezani s gravidnosti i laktacijom. To takoer zna i da je leptin potencijalno va~an indikator rane gravidnosti u krava pod uvjetom da u ispitivanje uklju imo joa neke imbenike koji e isklju iti utjecaj laktacije na razinu leptina u serumu. Na kraju smo analizirali i postojanje linearnih korelacija izmeu istra~ivanih parametara po skupinama. Pokazalo se da kod ~ivotinja koje su imale porast progesterona u serumu paralelno dolazi i do porasta leptina u serumu, ali pri tome ne postoje statisti ki zna ajne razlike izmeu skupina gravidnih, negravidnih i krava s ranom embrionalnom smrtnosti. `toviae, analiza linearnih korelacija u svim skupinama pokazala je da nema statisti ki zna ajnih korelacija izmeu pojedinih vrijednosti unutar skupine kao ni izmeu razli itih skupina. 6. ZAKLJU CI Na osnovi razine progesterona u serumu mogue je razlu iti gravidne od negravidnih krava ali nije mogue dijagnosticirati embrionalnu smrtnost. .Na osnovi razine trofoblast proteina u serumu (PAG-pregnancy associated glycoproteins) mogue je pouzdano dokazati postojanje ~ivog vitalnog embrija u maternici gravidne krave ve nakon 24.-tog dana gravidnosti. Na osnovi razine PAG mogue je sa sigurnoau razlikovati krave kod kojih je nastupila embrionalna smrtnost od krava koje nisu koncipirale ve nakon 24.-tog dana gravidnosti odnosno ako je embrij uginuo nakon 16.-tog dana gravidnosti . Na embrionalnu smrtnost ukazuje drasti an pad razine PAG u serumu majke i to 7,5-9 dana nakon uginua embrija, koliko iznosi i bioloaki polu~ivot PAG u prvom mjesecu gravidnosti Kriti no razdoblje za pre~ivljavanje embrija je izmeu 17. i 24. dana. U naaem istra~ivanju u tom periodu je embrij uginuo u 98% krava u skupini u kojoj je nastupila embrionalna smrtnost. Zbog utjecaja laktacije na razinu leptina u serumu ne mo~e se samo na osnovu tog parametra razlikovati negravidne, gravidne i krave u kojih je doalo do rane embrionalne smrtnosti, iako gravidnost najvjerojatnije utje e na razinu leptina u krvi krava. 7. SA}ETAK Za uspjeano praenje koncepcije i ranog tijeka gravidnosti neophodno je procijeniti sve faktore koji mogu utjecati na izostanak koncepcije ili ranu embrionalnu smrtnost. Svrha ovog rada bila je istra~iti ulogu trofoblast proteina (PAG) i leptina u fiziologiji rane gravidnosti u goveda kao i mogunost njihove primjene u dijagnostici rane embrionalne smrtnosti. Osobito je va~no razlikovati primarno neplodne jedinke od onih kod kojih je doalo do uginua embrija, kako bi se otkrili uzroci navedenih pojava a ~ivotinje nakon odgovarajueg tretmana ili lije enja im prije ponovno zabreale. U naaem istra~ivanju promatrane su 74 krave simentalske pasmine, u dobi 3-7 godina, koje su ve koriatene u reprodukciji i pri tome nisu imale nikakvih poteakoa s plodnoau. Sve su dr~ane na 5 obiteljskih gospodarstava u Meimurskoj ~upaniji, u uzgojima koji su brojili 8-20 krava. Krave su podijeljene u tri skupine: gravidne krave (n=32), krave sumnjive na ranu embrionalnu smrtnost (n=21) i negravidne krave (n=18). Na osnovi ultrazvu nih pretraga i laboratorijskog praenja razine leptina, progesterona i trofoblast proteina (PAG) u serumu od 12. do 45. dana nakon UO doali smo do slijedeih zaklju aka: Na osnovi razine progesterona u serumu mogue je razlu iti gravidne od negravidnih krava ali nije mogue dijagnosticirati embrionalnu smrtnost. Na temelju razine PAG mogue sa sigurnoau razlikovati krave kod kojih je nastupila embrionalna smrtnost nakon 16. dana gravidnosti od krava koje nisu koncipirale. Embrionalna smrtnost vidljiva je na osnovi drasti nog pada razine PAG u serumu majke i to 7,5-9 dana nakon uginua embrija, koliko iznosi i bioloaki polu~ivot PAG u prvom mjesecu gravidnosti. Praenjem razine PAG u serumu mogue je pouzdano dokazati postojanje ~ivog vitalnog embrija u maternici gravidne krave ve nakon 24. dana gravidnosti. Na osnovi razine leptina u serumu nije mogue razlikovati gravidne, negravidne i krave kod kojih je nastupila embrionalna smrtnost jer se utjecaj gravidnosti na razinu leptina u serumu preklapa s utjecajem laktacije na razinu leptina u serumu i nemogue je razlu iti razlike izmeu skupina iako rana gravidnost i sam embrij sigurno djeluju na razinu leptina u serumu. U naaem istra~ivanju je 98% krava u skupini u kojoj je nastupila embrionalna smrtnost, izgubilo embrij kad su bile gravidne 17-24 dana ato predstavlja kriti no razdoblje za pre~ivljavanje embrija. 8. ABSTRACT For succesful follow up of conception and early pregnancy flow it is necessary to evaluate all facts, potential causes of pregnancy failure or early embrionic mortality. The aim of this study was to investigate the role of pregnancy associated glycoproteins (PAG) and leptin in early pregnancy physiology in cows as well as possibility of using PAG and leptin as diagnostic tools in diagnosys of embrionic mortality. It is crucial in early pregnancy diagnosys to find unpregnant animals on time and return them in cyclicity and repeat AI. It is esspecially important to find and distinguish primary infertile cows from cows experienced embrionic mortality. In our research we used 74 cows of simmental breed aged 3-7 years, alrerady used regularly in reproduction and calved every year. All cows were kept on 5 family farms near akovec. All family farms consisted of 8-20 cows. We divided cows in 3 groups: pregnant cows (n=34), nonpregnant cows (n=18) and cows experienced embrionic mortality(n=21). According to ultrasound findings, and laboratory findings for progesteron, PAG and leptin in sera samples collected between 12. and 45. days after AI every 72 h, we concluded few things. FIrst of all, it is impossible to determine embrionic mortality on basis of progesterone profil, but it is easy to distinguish pregnant from nonpregnant cows, if cows are supposed to be pregnant more than 21. day. 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XII 1973. godine, gdje sam zavraila osnovnu i srednju akolu, zvanje veterinarski tehni ar. Veterinarski fakultet sam upisala u rujnu 1992. a diplomirala sam u velja i 1999. godine, s prosje nom ocjenom 4,16. Aktivno govorim engleski i njema ki jezik, za ato posjedujem meunarodni certifikat Goethe instituta. Tijekom studiranja sam uspjeano sudjelovala u dva meunarodna natjecanja u znanju stru nih predmeta za studente veterinarske medicine i osvojila dva priznanja:"Food for Though" za 1996. godinu, te "Best of Class" za 1997. godinu. Nadalje tokom studiranja, bila sam demonstrator na Zavodu za kemiju i biokemiju, Zavodu za anatomiju, histologiju i embriologiju te na Klinici za porodniatvo i reprodukciju. Poslijediplomski studij "Teriogenologija domaih sisavaca" sam upisala 2000. godine, a nakon odsluaana tri semestra, odobren mi je upis na doktorski studij bez izrade magistarskog rada. Po zavraetku studija krae vrijeme sam volontirala na Klinici za porodniatvo i reprodukciju, a potom sam u sije nju 2000., na istoj zaposlena kao stru ni suradnik, zamjena za mlaeg asistenta. Od rujna 2001. godine radim kao znanstveni novak na projektu MZO` 0053100 voditelja prof. dr. sc. Antuna Tomaakovia "Zna aj trofoblast proteina na pre~ivljavanje embrija papkara", a potom od lipnja 2003. kao stalni stru ni suradnik na Klinici za porodniatvo i reprodukciju. Tijekom 5 godina rada na Klinici za porodniatvo i reprodukciju, bila sam 9 puta na stru nim usavraavanjima u inozemstvu i to: - u lipnju 2002. na Institutu za biotehnologiju u Tullnu, Veterinarskog fakulteta u Be u - u rujnu 2002. Institut Ludwig-Maximillian, Veterinarskog fakulteta u Munchenu -u studenom 2002. CEEPUS stipendija, Klinika za porodniatvo, Veterinarskog fakulteta u Be u -u svibnju 2003. CEEPUS stipendija, Klinika za porodniatvo, Veterinarskog fakulteta u Be u -u lipnju 2003. pohaala sam Svjetsku akolu endokrinologije reprodukcije u pre~iva a za mlade znanstvenike, Budimpeata, Maarska -u studenom 2003. posjetila sam Zavod za fiziologiju reprodukcije, Veterinarskog fakulteta u Liege, Belgija - u rujnu 2004. u estvovala sam na ESDAR-u (radionice Europske udruge za reprodukciju u domaih ~ivotinja) Varaava, Poljska -u studenom 2004.i rujnu 2005. ponovno sam posjetila Zavod za fiziologiju reprodukcije, Veterinarskog fakulteta u Liege, Belgija u sklopu suradnje na znanstvenom projektu -u prosincu 2005. prisustvovala sam radionici ija je tematika bila reprodukcija u trkaih konja, odr~anoj u Newmarketu, Velika Britanija U okviru terenske nastave za studente organizirala sam i sprovodila kontrolu reprodukcije na ergeli engleskih punokrvnjaka Serdar u Blatu pored Zagreba tokom 2003, 2004 i 2005 godine. Autor i koautor sam na tridesetak znanstvenih radova, od ega je 5 objavljeno u tercijarnim publikacijama (Current Contents) a ostala u sekundarnim publikacijama te u zbornicima radova s brojnih domaih i meunarodnih kongresa. Koautorica sam na dvije skripte Primjenjena fiziologija reprodukcije u kobila i Reprodukcija u ovaca i koza.Odr~ala sam dva javna predavanja, jedno u Be u a drugo u Zagrebu, kao i pozvano predavanje na meunarodnom kongresu Clinica Veterinaria odr~anom na Ohridu. Takoer sam pisala i za asopis Veterinarstvo Nadalje, aktivno sam sudjelovala u pripremi IV Srednjeeuropskog bujatri kog Kongresa kao tehni ka tajnica. Tokom rada na klinici sudjelovala sam u voenju prakti ne i terenske nastave, suraivala sam na dva znanstvena projekta i sudjelovala i u svim ostalim aktivnostima koje Klinika obuhvaa. lan sam  Hrvatskog veterinarskog druatva ,  Hrvatske veterinarske komore , BEVA-e (British Equine Practitioners Veterinary Association) i  ESDAR -a (European Society for domestic animal reproduction)     PAGE  PAGE 8  :<@BH@H & F' & F%$a$<>BPH4\NRTJL "  " """ )))::@<B<f<h<== > >?? @καjx%h 'Uj`%h 'Uj%h 'Ujڇ%h 'Uj'f%h 'Uj<%h 'Uh 'H*^JaJ h 'H*^JaJ  jmh '^JaJ h '^JaJ h '6 jmh 'h ' hKg3CJ\ h 'CJ\j%h 'CJU\0HNPRT!!!\"#2#$%p&'() ) ))+,  & F)^`,-H/`0012j445x6699:::::b;;@<j<l<n< $ & F,da$$da$$da$  p#d & F(n<<"=X=Z==>>>>>>8>>X?Z??@,@.@0@BRF$a$d $ & F.da$ $hd^ha$ $ & F-da$$da$ @@@@@,@0@BTFFF\GGG HH2H4HXHZH~HHHHHHHHII4I6I\I^IIIIIIIII J"JHJJJlJnJJJJJJJKK6K8K^K`KKKKKKKKLL LHLJL󼴼h 'CJaJh 'CJ^JaJ h '5h '5CJ ^JaJ h '9\^JaJhKg35CJaJh '5CJaJh 'jV&h 'UGRFFFZG\GtGGGGG $$Ifa$$a$ GGGGHH HH????? $$Ifa$kd%&$$IfTl<rg c'T'T'T'T0#644 laT HHHH"H*H2HH????? 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