Vaňo, M. - Bučko, O. - Kováč, Ľ. - Dvořák, J. - Matoušek, V. - Čopík, A. Slovenská poľnohospodárska univerzita Nitra
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1 ASOCIÁCIA ALEL GÉNU FOLIKULOSTIMULAČNÉHO HORMÓNU (FSHB) KU ZNAKOM REPRODUKCIE ASSOCIATION OF GENE ALLELES OF FOLLICULAR STIMULATIVE HORMONE (FSHB) TO REPRODUCTIVE SINGNS Vaňo, M. - Bučko, O. - Kováč, Ľ. - Dvořák, J. - Matoušek, V. - Čopík, A. Slovenská poľnohospodárska univerzita Nitra The aim of our thesis was to pursue the gene FSHB with respect to single alleles and their influence on chosen reproduction parameters of sows. By the frequency evaluation of the allele by gene FSHB we found out the frequency of the allele A 0,05 and allele B 0,95. The genotype frequences were: AA 0 %, AB 9,30 % and BB 90,7 %. By the analysing of the single genotypes FSHB we found out the highly statistical evidentiary differences: in the number of live-born pigs between the genotypes BB 11,14 and AB 9,38 and the statistical evidentiary differences of brought forward pigs count AB 1,12 heads and BB 0,06 heads and of the dead-born pigs AB 0,38 heads and BB 1,47 heads. We did not find out the significant influence of marker FSHB on the pursued reproduction parameters. At average for lifelong sow efficiency the differences of the pursued reproduction parameters were not significantly influenced by marker FSHB. The analyses of the sow genotypes, marker genotypes and their interactions, correlations, multilinear regressions and the contingent tabelles showed on the necessary of more detailed solutions by problem investigation in our thesis by the planned animal scientific-genetic experiments, on the basis which it could be possible to obtain the knowledges applicable in the sow selection on the influencing of the level by the sow reproduction parameters. Key words: marker, reproduction, sow, genotype Cieľom našej práce bolo sledovať gén FSHB s ohľadom na jednotlivé alely a ich vplyv na vybrané reprodukčné parametre prasníc. Pri vyhodnocovaní frekvencie alel génu FSHB sme zistili frekvenciu alely A 0,05 a alely B 0,95. Frekvencie genotypov boli: AA 0 %, AB 9,30 % a BB 90,7 %. Pri analýze jednotlivých genotypov FSHB na prvom vrhu prasníc sme zistili vysoko štatisticky preukazné rozdiely: počtu živonarodených prasiat medzi genotypmi BB 11,14 a AB 9,38 a štatisticky preukazné rozdiely počtu presunutých prasiat AB 1,12 ks a BB 0,06 ks a uhynutých prasiat AB 0,38 ks a BB 1,47 ks. Na druhom vrhu prasníc sme nezistili signifikantný vplyv markeru FSHB na sledované reprodukčné ukazovatele. V priemere za celoživotnú úžitkovosť prasníc rozdiely sledovaných reprodukčných ukazovateľov neboli signifikantne ovplyvnené markerom FSHB. Analýzy genotypov prasníc, genotypov markerov ich interakcií, korelácií, viacnásobných lineárnych regresií a kontigenčných tabuliek poukázali na potrebu detailnejšieho skúmania problematiky študovanej v našej práci pri plánovaných zootechnicko-genetických experimentoch, na základe ktorých by bolo možné získať poznatky aplikovateľné v selekcii prasníc na ovplyvnenie úrovne reprodukčných ukazovateľov prasníc. Kľučové slová: marker, reprodukcia, prasnica, genotyp Úvod: 492
2 Rozvoj molekulárno-genetických metód umožnil zahájenie analýz genómu hospodárskych zvierat. Rastie rad poznatkov o nepredstaviteľnej premenlivosti DNA v kódujúcich a nekódujúcich oblastiach a o riadení génovej expresie. Využitie metód molekulárnej genetiky pri hospodárskych zvieratách sa sústredilo na priame štúdium kandidátnych génov, stanovené na základe biochemických vlastností génového produktu a jeho úlohy v metabolizme (funkčné kandidátne gény), alebo na základe komparácie s inými živočíšnymi druhmi (pozičné kandidátne gény). Cieľom práce bolo sledovať gén FSHB (gén folikulostimulačného hormónu) s ohľadom na jednotlivé alely, stanoviť genotypy vybraného génu metódami molekulárnej genetiky PCR - RFLP a vyhodnotiť vzťah uvedeného génu k sledovaným ukazovateľom reprodukcie. Literárny prehľad: Gén FSHB (follicular-stimulating hormone beta subunit) bol lokalizovaný na chromozóme 2 (Ellegreen et al., 1994). Nukleotidová sekvencia génu FSHB vyštepuje veľký počet kópií. Polymorfizmus tohto génu u ošípaných bol identifikovaný reštrikčným enzýmom HaeIII. V roku 1994 Rohrer et al., (1994) vplyv polymorfizmu génu FSHB na reprodukčné vlastnosti nepreukázali. Pri štúdii ktorú uskutočňovali Linville et al., (2001) boli porovnávané tri línie. Rozdiely medzi líniou selektovanou na ovuláciu (IOL) a líniou kontrolnou (C) bolo 6,1 ovárií a 4,7 vyvinutých prasiatok na vrh. Medzi líniou selektovanou na veľkosť vrhu (COL) a kontrolnou (C) boli rozdiely 2,2 ovárií a 2,9 vyvinutých prasiatok na vrh. Diferencie boli zistené i vo frekvencii alel génu FSHB. Pri línii IOL bola frekvencia alely B o 0,35 vyššia ako pri línii C (0,97. resp. 0,62) a medzi líniami COL a C bol tento rozdiel 0,15 (0,77 resp. 0,62). Nenašli sa však signifikantné rozdiely medzi polymorfizmom markeru a fenotypom reprodukčných vlastností. Doterajšie štúdie v Českej republike ukazujú, že prasnice kombinovaného genotypu BBBB génov ESR a FSHB všeobecne rodia o 1,85-3,01 viac všetkých narodených prasiat a o 2,0-3,0 viac živonarodených prasiat než prasnice kombinovaného genotypu ABAA (Putnová et al., 2002). Štruktúru génu popísal Hirai et al. (1990) a Rohrer et al. (1994) identifikoval v géne HaeIII PCR-RFLP. Li et al. (1998) našli asociáciu FSHB génu s veľkosťou vrhu. Analýzou čínskych a západných plemien (landras, yorkshire, hybridy plemien yorkshire a Erhulian) našli signifikantný vplyv alely B na veľkosť vrhu. Priemerný efekt genotypov BB na veľkosť vrhu bol o 2,53 a 2,12 prasiat na vrh viac na celkový počet narodených a živonarodených prasiat u prvých vrhov. Pre druhý, tretí a štvrtý vrh sa efekt genotypu BB znižuje, ale aj napriek tomu homozygoti pre priaznivú alelu vykazujú efekt o 1,5 prasiat na vrh viac než prasnice opačného genotypu. Pri všetkých vrhoch sa objavil pozitívny efekt a pohyboval sa okolo jedného prasaťa. Huang et al. (2000) zistili u prasníc genotypu BB plemena yorkshire, durok, landras, Erhulian, Xiang a Wuzhishan o 0,55-2,21 živonarodeného prasaťa na vrh viac ako u prasníc genotypu AA. Kossakowska et al. (2000) nenašli u syntetickej línie 990 preukazné diferencie medzi genotypmi (frekvencie alely B 0,892, len 3 prasnice genotypu AA). Li et al. (2000) odhalili prítomnosť novej detekovanej alely B len u plemena meishan a ďalších 5 čínskych plemien (Erhulian, Fengjing, Hainan, Min a Xiang), alelu nemal americký landras, Large White a ošípané Berkshire. Chen et al. (2001) sledovali ako kombináciu genotypov pre gény ESR a FSHB, medzi ktorými nie sú žiadne genetické interakcie, ovplyvní plodnosť prasníc (n = 269). Prišli k záveru, že prasnice kombinovaného genotypu BBBB majú o 1,85-3,01 všetkých narodených a o 2,0-3,0 živonarodených prasiat viac než prasnice genotypu ABAA. Gén FSHB vykazuje extrémne asymetrickú distribúciu 493
3 genotypov. Pri komerčných plemenách bol zistený veľký výskyt genotypu BB, zatiaľ čo u čínskych plemien sa vyskytuje len zriedka. Materiál a metodika Hodnotená bola reprodukčná úžitkovosť vybraných prasníc plemená biela mäsová (BM) a dvojplemenných, resp. trojplemenných krížencov na ktorých sa podieľali plemená landras (L), durok (D) a holandský hybrid dalland (DA). Hodnotená bola reprodukčná úžitkovosť u nasledovných genotypov prasníc: BM biela mäsová 26 ks BML biela mäsová x landras 23 ks (BML)L (biela mäsová x landras) x landras 10 ks (BML)BM (biela mäsová x landras) x biela mäsová 5 ks (BMDA)L (biela mäsová x daland) x landras 9 ks (BMDA)BM (biela mäsová x daland) x biela mäsová 4 ks (BMD)L (biela mäsová x durok) x landras 7 ks (BMD)BM (biela mäsová x durok) x biela mäsová 2 ks Spolu 86 ks Sledované reprodukčné ukazovatele: - počet živonarodených prasiat vo vrhu v ks (ŽP) - počet mŕtvonarodených prasiat vo vrhu v ks (MP) - počet anomálnych prasiat v ks (AP) - počet prasiat s anomálnym intrauterinným vývojom - počet presunutých prasiat v ks (PP) - počet priložených resp. preložených prasiat medzi jednotlivými vrhmi prasníc - počet uhynutých prasiat vo vrhu do odstavu v ks (UP) - počet odstavených prasiat v ks (OP) - počet mumifikovaných prasiat v ks (MUP) - počet prebehov prasníc do úspešnej inseminácie (PPP) - počet opakovaných estrálnych cyklov prasníc po úspešnú insemináciu - počet neproduktívnych dní do 1. pripustenia prasníc (NDP1) - interval od odstavu po prvú insemináciu prasníc (v dňoch) - počet neproduktívnych dní od 1. pripustenia po úspešné pripustenie prasníc (NDPU) - interval od prvej inseminácie po oplodnenie prasníc (v dňoch) Výsledky a diskusia V tabuľke I. uvádzame výskyt frekvencie genotypov a alel sledovaného markera spolu za celú populáciu prasníc. Pri vyhodnocovaní frekvencie alel génu FSHB sme zistili frekvenciu alely A 0,05 a alely B 0,95. Frekvencie genotypov boli: AA 0 %, AB 9,30 % a BB 90,7 %. Tab. I.: Frekvencie genotypov alel sledovaných markerov spolu za celú populáciu prasníc Marker P H Q n p q FSHB AA AB BB A B ,05 0,95 0 % 9,3 % 90,7 % 100 % V našej práci sme overovali vplyv polymorfizmu génu folikuly stimulujúceho hormónu (FSHB) na sledované reprodukčné parametre prasníc uvedené v tabuľke II.. 494
4 Tab. II.: Dosiahnuté parametre reprodukcie pri jednotlivých genotypoch FSHB na 1., 2. vrhu a v priemere za celoživotnú úžitkovosť prasníc Ukazovateľ ŽP MP AP PP UP Poradie Genotypy FSHB Genotypy FSHB Ukazovateľ vrhu n AB n BB n AB n BB 1. vrh 8 9, ,14 8 9, ,67 2. vrh 8 11, ,17 OP 8 10, ,88 celož. úžit. 8 11, , , ,84 1. vrh 8 1, ,60 8 0, ,08 2. vrh 8 0, ,47 MUP 8 0, ,03 celož. úžit. 8 0, ,69 8 0, ,09 1. vrh 8 0, ,76 8 0, ,26 2. vrh 8 0, ,59 PPP 8 0, ,26 celož. úžit. 8 0, ,84 8 0, ,16 1. vrh 8 1, ,06 8 0, ,55 2. vrh 8-0, ,78 NDP1 8 15, ,94 celož. úžit. 8-0, ,85 8 6, ,87 1. vrh 8 0, ,47 8 0, ,18 2. vrh 8 1, ,29 NDPU 8 0, ,08 celož. úžit. 8 1, ,14 8 1, ,58 Pri analýze jednotlivých genotypov FSHB sme na prvom vrhu prasníc zistili väčší počet živonarodených prasiat pri genotype BB 11,14 ± 0,21 a menší počet pri genotype AB 9,38 ± 0,63. Prítomnosť genotypu AA sme DNA analýzou v testovanej skupine prasníc nezistili. Rozdiely medzi jednotlivými genotypmi FSHB boli vysoko štatisticky preukazné. Naše výsledky potvrdzujú tvrdenia doterajších štúdií v Českej republike, ktoré uvádza tiež Putnová et al. (2002). Autorka poukazuje na to, že prasnice kombinovaného genotypu BBBB génov ESR a FSHB všeobecne rodia o 1,85-3,01 ks všetkých narodených prasiat a o 2,0-3,0 ks živo narodených prasiatok viac, než prasnice kombinovaného genotypu ABAA. Naše výsledky sa zhodujú aj s výsledkami Královej (2003), ktorá zistila štatisticky preukazný vplyv génu FSHB na počet živonarodených prasiat na prvom vrhu prasníc superplodnej línie v prospech BB genotypu (AA 11,65; AB 11,82; BB 12,98). Zistili sme štatisticky významné rozdiely premenlivosti počtu presunutých prasiat spôsobené markerom FSHB. Pri genotype AB sme zistili presun 1,12 ks a pri genotype BB 0,06 ks prasiat. Pri celkovom hodnotení v rámci genotypov FSHB sme zistili štatisticky významný vplyv markeru FSHB na počet uhynutých prasiat: pri genotype AB 0,38 ks a pri genotype BB 1,47 ks. Pri ostatných sledovaných reprodukčných ukazovateľoch sme nepotvrdili vplyv genotypu FSHB na ich premenlivosť. Zistili sme štatisticky vysoko významný vplyv genotypu prasnice na počet mumifikovaných prasiat. Rozdiely premenlivosti hodnotených ukazovateľov spôsobené interakciou genotyp prasnice a genotyp FSHB boli štatisticky nepreukazné. Na druhom vrhu prasníc sme nezistili štatisticky preukazný vplyv markeru FSHB na žiadny hodnotený reprodukčný ukazovateľ. Zistili sme štatisticky preukazné rozdiely počtu mŕtvonarodených a anomálnych prasiat a štatisticky vysoko preukazné rozdiely počtu mumifikovaných prasiat spôsobené genotypom prasnice. Rozdiely počtu anomálnych prasiat a počtu neproduktívnych dní do úspešnej inseminácie spôsobené interakciou genotyp prasnice a genotyp FSHB boli štatisticky preukazné a vysoko preukazné v počte mumifikovaných prasiat. Nepotvrdili sme výsledky Královej (2003), ktorá na druhom a ďalších vrhoch prasníc zistila štatisticky preukazný vplyv markeru FSHB na počet živonarodených prasiat medzi genotypmi BB a AB (AA 14,05; AB 12,85; BB 13,85). Potvrdili sme však tvrdenia, že genotyp FSHB štatisticky preukazne neovplyvňuje počet odchovaných prasiat, rovnako ako to uvádza autorka vo svojej práci (AA 12,76; AB 11,53; BB 11,97). 495
5 Rozdiely sledovaných reprodukčných ukazovateľov v priemere za celoživotnú úžitkovosť prasníc neboli štatisticky preukazne ovplyvnené genotypom FSHB, genotypom prasníc ani interakciou genotyp prasnice a marker ESR. Analýzou rozptylu hodnotených ukazovateľov sme zistili štatisticky významnú regresiu priemerného počtu uhynutých prasiat v závislosti od počtu vrhov prasníc 0,106 ks a vysoko štatisticky preukaznú regresiu priemerného počtu odstavených prasiat od počtu vrhov prasníc -0,247 ks. Záver: Z rozboru zistených výsledkov vyplýva význam hodnotenia reprodukčných ukazovateľov prasníc nielen z hľadiska ich genetického založenia, ale tiež z hľadiska markerov, ktoré ich determinujú pre selekciu prasníc. Je možné odporučiť ďalšie testovanie génu FSHB s ohľadom na jeho genotypy hlavne genotyp BB. Pre praktické využitie našich výsledkov, ako aj iných podobných štúdií, je potrebné inštalovať vhodné optimálne plány zootechnicko-genetických experimentov tak, aby sme pri vynaložených nákladoch získali optimálne výsledky realizovateľné v selekcii prasníc. Zistené výsledky sú tiež podnetom pre ďalšie testovanie a analyzovanie všetkých faktorov ovplyvňujúcich reprodukciu prasníc, vrátane novoobjavených kandidátnych génov. Zoznam použitej literatúry: 1. ELLEGREN, H. - CHOWDHARY, B. P. - JOHANSSON, M. - MARKLUND, L. - FREDHOLM, M. - GUSTAVSSON, I. - ANDERSSON, L.: A primary linkage map of the porcine genome reveals a low rate of genetic recombination. Genetics 137, , HIRAI, T. TAKIKAWA, H. KATO, Y.: The gene for the beta subunit of porcine FSH: absence of consus oestrogen-responsive element and presence of retroposons. Journal of Molecular Endocrinology 5, , HUANG, L. S. CHEN, K. F. LI, N. REN, J. DING, N. S. MEYER, J. N. BESENKO, S. P. GAO, J. LUO, M.: Genetic variations at the type I and type II markers loci in different Chinese local and Western commercial pig breeds. 27 th International conference on Animal Genetics, July 22-26, University of Minnessota, Minneapolis, 66, CHEN, K. LI, N. - HUANG, L. ZHANG, Q. ZHANG, J. SUN, S. LUO, M. WU, C.: The combined genotypes effect of ESR and FSHB genes on litter size traits in five different pig breeds. Chinese Science Bulletin 46, , KOSSAKOWSKA, A. KAMYCZEK, M. KURYL, J. PERZCHALA, M. CIESLAK, D.: Polymorphism in the FSHB and OPN gene and their association with reproductive traits in synthetic pig line th International Conference on Animal Genetics, July 22-26, University of Minnesota, Minneapolis, 72, KRÁLOVÁ, P.: Šlechtení superplodných linií mateřských plemen prasat. Doktorská disertační práce. Jihočeská univerzita v Českých Budějovicích, Zemědelská fakulta, s. 184, LI, M. D. ZHAO, Y. F. XIAO, L. ZHANG, F. J. CHEN, Y. Z. DAI, R. J. ZHANG, J. S. SHEN, S. Q. CHEN, Y. F. WU, C. X.: candidate gene approach for identification of genetic loci controlling litter size in swine. Proceedings of the 6 th World Congress on Genetics Applied to Livestock Production 26, , LI, M. D. ROHRER, G. A. WISE, T. H. FORD, J. J.: Identification and characterization of a new allele for beta subunit of follicle-stimulating hormone in Chinese pig breeds. Animal Genetics 31, 28-30, LINVILLE, R. C. - POMP, D. - JOHNSON, R. K. - ROTHSCHILD, M. F.: Candidate gene analysis for loci affecting litter size an avolution rate in swine. J. Anim. Sci., 79: 60-67, PUNTOVÁ, L. - KOLAŘÍKOVÁ, O. - DVOŘÁK, J.: Association of genetic markers for litter size in materiel pig lines. In: Current problems of breeding, health and production of pigs, České Budějovice: 30 s, ROHRER, G. A. - ALEXANDER, L. J. - KEELE, J. W. - SMITH, T. P. - BEATTIE, C. W.: A microsatellite linkage map of the porcine genome. Genetics 136, ,
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