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Disertační Práce MASARYKOVA UNIVERZITA V BRNĚ Přírodovědeckáfakulta DISERTAČNÍ PRÁCE Mgr.JanMendel Brno,2007 MASARYKOVA UNIVERZITA V BRNĚ Přírodovědeckáfakulta Ústavexperimentálníbiologie Ústavbiologie obratlovcůAVČR,v.v.i. Mgr. Jan Mendel Využití mtDNA pro charakterizaci populačních struktur druhů rodů Gobio a Romanogobio Disertační práce Školitel: RNDr. Věra Lusková, Ph.D. Brno, 2007 1 Bibliografickáidentifikace Jménoa příjmeníautora:Mgr.JanMendel Názevdisertační práce: VyužitímtDNA procharakterizaci populačníchstrukturdruhůrodů Gobio a Romanogobio . Názevdisertační práceanglicky:TheusingofmtDNA for characteristicsof population structuresofspeciesfromthegenera Gobio and Romanogobio . Studijní program:Biologie Studijníobor(směr),kombinaceoborů:Obecnáamolekulárnígenetika Školitel:RNDr.VěraLusková,Ph.D. Rokobhajoby:2007 Klíčováslovavčeštině: Gobio , Romanogobio , taxonomie, systematika, fylogeneze, fylogeografie, RAPD, sekvencování, mtDNA, nDNA, control region, cytochrom b, S7 r-protein, intron, S7indel diagnostika, hybridizace, vnitrodruhováamezidruhovádiverzita Klíčováslovavangličtině: Gobio , Romanogobio , taxonomy, systematics, phylogeny, phylogeography, RAPD, sequencing, mtDNA, nDNA, control region, cytochrome b, S7 r-protein,intron, S7indel diagnostics, hybridization, intraspecificandinterspecificdiversity 2 ©JanMendel,MasarykovauniverzitavBrně,2007 3 PODĚKOVÁNÍ Tuto disertační práci věnuji své ženě Denisce jako dík za vždy teplou večeři, hřejivé obětí a za oporu,kterou promě vmém životě trvale představuje.Malémuještě nenarozenému Samuelovi,kterýbyl vzávěrumým největším hnacím motorem.A také celé své rodině,která měneustále podporovalaa podporuje. Rád bych poděkoval Věře Luskové, Stanislavu Luskovi a ostatním kolegům z ichtyologického oddělení Ústavu biologie obratlovců, v.v.i., Lukáši Vetešníkovi, Karlu Halačkovi,IvuPapouškoviaJanuKociánovi,beznichž bytatoprácenemohlavzniknout. Rádbychtaké poděkoval domácím i zahraničním kolegům JánuKoščovi,Ekaterine D. Vasil’eve,JohnuTaggartovi,AlexandruRuchinovi,Guler Ekmekci,FüsunErk’akan,Cevher S. Özeren, Radku Šandovi, Andrei N. Pashkovovi, Sergei I. Reshetnikovovi a mnohým dalším,kteří mi poskytli svévzorkya přispělisvými podnětykřešení této problematiky. Tato dizertační práce byla řešena vrámci výzkumných projektů: IAA6045005 „Charakteristiky a genetické rozdílnosti populací a druhů „hrouzka“ rodů Gobio a Romanogobio –příspěvek kpoznání biodiverzity“ Grantové agentury AV ČR, IPS5045111 „Molecular and other genetic markers applied in the conservation of populations of endangered, rare andvanishingfish species inthe CzechRepublic“ Grantové agenturyAV ČR a VaV/SM/6/3/05„Genetická diverzita ohroženýchdruhůryb –nezbytnýzákladefektivní ochranybiodiverzity“ MinisterstvaživotníhoprostředíČR. 4 Prohlašuji, že tuto práci jsem vypracoval samostatně. Veškerou literaturu a ostatní prameny, znichž jsem při přípravě práce čerpal, řádně cituji a uvádím vseznamu použité literatury. 5 Abstrakt VyužitímtDNA procharakterizacipopulačníchstrukturdruhůrodů Gobio a Romanogobio Taxonomie a systematika druhů rodů Gobio a Romanogobio je vsoučasné době velmi nepřehledná.O validitě řadydruhů,poddruhůa forem,jež bylypopsányvminulosti,se vedou diskuse. Současná systematika obou rodů je založena především na morfologických charakteristikácha nebyla dosudpotvrzena molekulárními analýzami.Tatostudie se zabývá fylogenetickými vztahy 28 druhů obou rodů veuroasijském kontextu. Genetická diverzita hrouzků byla zkoumána technikami RAPD a přímým sekvencováním control regionu, cytochromu b a intronu S7 r-proteinu. Molekulární analýzy potvrdily rodovou diferenciaci původního rodu Gobio na dva rody Gobio a Romanogobio . Bylo prokázáno, že území ČR osídlují druhy Gobio gobio , Gobio obtusirostris , Gobio carpathicus (ojediněle), Romanogobio banaticus , Romanogobio vladykovi , Romanogobio belingi a druh označený termínem „species-in-Waiting“ Romanogobio carpathorossicus . Na území SR se vyskytuje vedle tří výše jmenovanýchdruhůrodu Gobio i čtvrtýdruh pracovně označený Gobio sp.1. Rod Romanogobio má obdobnou druhovou skladbu jako v ČR vyjma druhu Romanogobio belingi .Na slovenském území byl navíc potvrzenvýskyt druhu Romanogobiofriči .Výsledky ukázaly, že řekyČernéhomoře vykazují větší druhovoupestrost rodu Gobio než řeky tekoucí do Severního a Baltského moře. Celkem bylo zachyceno 16 hybridních událostí, které se týkaly obourodů.Na území SR byla vymezena hybridní zóna voblasti severních přítokůřeky Tisy.Byla posouzena genetická odlišnost populací na území ČR a SR,především ukriticky ohroženýchdruhůrodu Romanogobio .U šesti populací druhu Gobiogobio a dvoupopulací Gobio obtusirostris byla zjištěna statisticky významná mezipopulační diverzita. Bylo odhaleno celkem 40 populačně diagnostických RAPD markerů. Byla nalezena jednoduchá rozlišovací metoda nazvaná „S7indel diagnostika“, která se osvědčila jako nástroj rychlé identifikace druhů rodu Gobio . Bylo navrženo rozšíření subjektu ochrany o „ochranu genetické diverzity“ jakovsoučasnosti nejsnáze uplatnitelnýpostup při ochraně biodiverzity vČR. 6 DissertationAbstract TheusingofmtDNAforcharacteristicsofpopulationstructuresof species from the genera Gobio and Romanogobio Taxonomy andsystematics of species of the genera Gobio and Romanogobio is rather confusedat present.Validityof a number of species,subspecies andforms describedinthe past is under discussion.The present systematics of both the genera is based especially on morphological characteristics and has not been confirmed by molecular analyses yet. The studydeals Withphylogenetic relationships of 28species of boththe genera inthe EuroAsian context. Genetic diversityof gudgeons Was investigatedusingthe RAPD technique anddirect sequencingof the control region,cytochrome b andthe intronof the S7 r-protein.Molecular analyses have confirmedgeneric differentiation of the original genus Gobio intotWogenera Gobio and Romanogobio .It has beendemonstratedthat the territoryof the CzechRepublic is inhabited by the species Gobio gobio , Gobio obtusirostris , Gobio carpathicus (only one specimen found), Romanogobio banaticus , Romanogobio vladykovi , Romanogobio belingi and by the species Romanogobio carpathorossicus Which is denoted as a “species-in- Waiting”. Concerning the territory of the Slovak Republic, in addition to the three above mentionedspecies of the genus Gobio alsoa fourthone occurs there Whichis provisionally called Gobio sp. 1. The genus Romanogobio has a similar species composition as in CR except for Romanogobio belingi . In addition, the presence of Romanogobio friči Was confirmed in the territory of Slovakia. The results have pointed out, that the rivers of the BlackSea show greater species diversityof the genus Gobio thanthe rivers flowingintothe NorthandBaltic Seas. Altogether 16hybridizationevents concerning boththe genera Were recorded.In the territory of the SlovakRepublic, a hybrid zone Was delineated in the area of notherntributaries of the Tisa River.Genetic difference of the populations inthe territories of CR and SR Was examined, especially in the critically endangered species of the genus Romanogobio .Statisticallysignificant inter-populationdiversityWas foundinsixpopulations of Gobio gobio and tWo populations of Gobio obtusirostris . Altogether 40 population diagnostic RAPD markes Were identified. A simple differentiation method called “S7indel diagnostics” Was found; it provedtobe a useful tool for quickidentificationof the species of the genus Gobio . Broadening of the subject of conservation With “genetic diversity conservation” Was suggestedas the easiest procedure inbiodiversityconservationinCR at present. 7 OBSAH 1 Úvod .................................................................................................................................10 2 Přehledsoučasnéhostavuřešenéproblematiky ..........................................................14 2.1 Diverzitadruhováavnitrodruhová–obecnýpohled ...............................................14 2.2 Taxonomie zkoumanéhoobjektu .............................................................................17 2.3 Použiténázvosloví....................................................................................................19 2.4 Hrouzekobecný( Gobio gobio Linnaeus,1758) ......................................................20 2.4.1 RozšířenívČRaSR.........................................................................................21 2.4.2 Stupeňohroženíaochrana................................................................................21 2.5 HrouzekKesslerův( Romanogobiokesslerii Dybowski,1862) ...............................22 2.5.1 RozšířenívČRaSR.........................................................................................23 2.5.2 Stupeňohroženíaochrana................................................................................24 2.6 Hrouzekběloploutvý( Romanogobioalbipinnatus Lukasch,1933).........................26 2.6.1 RozšířenívČRaSR.........................................................................................27 2.6.2 Stupeňohroženíaochrana................................................................................30 2.7 Hrouzekdlouhovousý( Romanogobiouranoscopus Agassiz,1828) .......................32 2.7.1 RozšířenívČRaSR.........................................................................................32 2.7.2 Stupeňohroženíaochrana................................................................................33 2.8 Hybridizaceaintrogrese...........................................................................................35
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