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Uticaj Biljke Domaćina Na Kriptičnu Diferencijaciju Populacija Vektora

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Uticaj Biljke Domaćina Na Kriptičnu Diferencijaciju Populacija Vektora UNIVERZITET U BEOGRADU BIOLOŠKI FAKULTET Andrea M. Kosovac Uticaj biljke domaćina na kriptičnu diferencijaciju populacija vektora Hyalesthes obsoletus Signoret, 1865 (Hemiptera: Cixiidae) i epidemiološke puteve prenošenja ‘Candidatus Phytoplasma solani’ doktorska disertacija Beograd, 2018 UNIVERSITY OF BELGRADE FACULTY OF BIOLOGY Andrea M. Kosovac The influence of host-plant use on cryptic differentiation of vector Hyalesthes obsoletus Signoret, 1865 (Hemiptera: Cixiidae) and on epidemiological transmission routes of ‘Candidatus Phytoplasma solani’ Doctoral Dissertation Belgrade, 2018 Mentori: _________________________________________ prof. dr Željko Tomanović, redovni profesor Biološki fakultet Univerzitet u Beogradu _________________________________________ dr Jelena Jović sredinu , viša naučna saradnicaBeograd Institut za zaštitu bilja i životnu Članovi komisije: _________________________________________ dr Ivo Toševski , naučni savetnikBeograd Institut za zaštitu bilja i životnu sredinu Datum odbrane: _____________ Doktorska disertacija urađena je u Institutu za zaštitu bilja i životnu sredinu, Odseku za štetočine bilja u Zemunu, u okviru projekta Ministarstva prosvete, nauke i tehnološkog razvoja br. III43001 - „Agrobiodiverzitet i korišćenje zemljišta u Srbiji: integrisana procena biodiverziteta ključnih grupa artropoda i biljnih patogena“ (rukovodilac prof. dr Željko Tomanović). Deo populaciono-genetičkih istraživanja je urađen na Katedri za Zoologiju, Odseka za Ekologiju, Johan Gutenberg Univerziteta u Majncu u saradnji sa profesorom dr Jesom Johannesenom. Morfološka identifikacija H. obsoletus primeraka asociranih sa V. agnus- castus i C. foetida je urađena u saradnji sa profesorkom dr Hannelore Hoch iz Prirodnjačkog muzeja u Berlinu. *** Zahvaljujem mentorki dr Jeleni Jović na prenetom znanju i mentoru profesoru dr Željku Tomanoviću na savetima i podršci. Hvala kolegama Odseka za štetočine bilja, Instituta za zaštitu bilja i životnu sredinu u Beogradu: dr Ivu Toševskom, dr Milani Mitrović, dr Tatjani Cvrković, dr Oliveru Krstiću i Miljani Jakovljević na pomoći u istraživanjima, prijateljskim razgovorima i savetima. Hvala dr Sanji Radonjić i dr Snježani Hrnčić sa Biotehničkog fakulteta, Univerziteta u Podgorici, na pomoći pri sakupljanju materijala. Zahvaljujem profesorki dr Hannelore Hoch na muzejskom paratipskom primerku vrste Hyalesthes thracicus ustupljenom iz lične entomološke kolekcije. Zahvalnost dugujem profesoru dr Jesu Johannesenu za preneto znanje i prijateljsku podršku čime ga smatram neformalnim mentorom. Posebno se zahvaljujem mojim roditeljima na nesebičnoj podršci i razmevanju. UticajHyalesthes biljke domaćina obsoletus na kriptičnuSignoret, 1865diferencijaciju populacija vektora prenošenja ‘Candidatus(Hemiptera: Cixiidae) i epidemiološke puteve Phytoplasma solani’ SAŽETAK Hyalesthes: obsoletus, vektor biljnog patogena 'Candidatus (Ca solani', polifagn .) Phytoplasma Evropi ismatra Mediteranu se om štetočin sprovedenom od ekonomskog značaja u centralnoj . Istraživanjima im u jugoistočnoj 'Ca Evropi,. P. solani' koja- gravitirauslovljenih centru biljnih arela bolesti, vrste i gde je zabeležen značajan broj utvrđenoConvolvulus je postojanje arvensis (četiriCa , Urtica asocijacije dioica vektora(Ud , Vitex sa biljkamaagnus-castus dualnim (Vac domaćinima: i Crepis foetida (Cf . ) - ) ) ) Populaciono genetičkim analizamana mitohondrijskih i nuklearnih markera (mikrosatelita)sintopsk i geografski utvrđena udaljenihje kriptič H. diferencijacijaobsoletus simpatričkih (uključujući e) Ca-Ud populacija Vac i naCf .tri Pored filogenetičke jasne linije uslovljene na biljkom domaćinom: gen (1) (1.1%, -(2) (3) diferencijacijeizdvajanje mitohondrijskim ima klastera 1.5%) i mikrosatelita koji (pripadnost potvrđuju >90% populacija u jedan od tri premaH. obsoletusbiljci domaćinu sensu lato ), podršku za izdizanjeregistrovana genetičke divergencijeu odnosu na najsrodnijuna nivo kriptične vrstu - specijacijeHyalesthes dajethracicus (2.1% - distanca3.3% . morfološki H. obsoletus sensu lato ) Posledice specijalizacije i diferencijacije'Ca. prema biljkama domaćinima na epidemiologijudetektovanih P. solani' procenjenein situ su multigenskom karakterizacijom izolata fitoplazme vinograda(1) i (2 u prirodnim epidemiološkimunakrsne in sistemima biljaka mediteranskih h ) eksperimentima. okulacije putem prirodno 'inficiraniCa. P. solani'populacija na tuf-a/- abvektora i tuf-b Potvrđena je osnovnaUd i Caepidemiološka uz ukrštanje divergencijasa Vac- tuf- cikluse vezane za tuf- uzrokovanom b epidemiologijom i utvrđenalinija vektora, kompleksnost H. obsoletus ex bCf ,epidemiologije kao u kojoj učestvuje -i C.treća foetida genetička. i sama biljka dualni domaćin KLJUČNE REČI Hyalesthes obsoletus, , : , e biljka domaćin populaciona genetika, NAUČNAkriptična evolucija,OBLAST: Biologijastolbur fitoplazma pidemiologija, multigenska karakterizacija UŽA NAUČNA OBLAST: UDK: [[577.832+595.753]Entomologija] 579.887 :576.89 : (043.3) -plant use of TheHyalesthes influence obsoletus of host Signoret, 1865on cryptic differentiation andvector on (Hemiptera: of ‘Candidatus Cixiidae) epidemiological transmission routes Phytoplasma solani’ ABSTRACT: Hyalesthes obsoletus The cixiid planthopperCandidatus (Ca. is generally considered as a polyphagous vector of ' ) P. solani' associated with numerous wild and cultivated plants in central Europe and Mediterranean.H. obsoletu sResearch conducted in southeastern'Ca. P. solani' Europe,- the distribution centre of and the area of- plantmost inflictedConvolvulus crop diseases,arvensis ( Capoints, Urtica toward dioica specific (Ud , Vitex dual agnus host-castus associations(Vac and Crepis with: foetida (Cf -) ) ) ). Population genetic analysis of mitochondrial and nuclear genetic markers (microsatellites) have revealed cryptic differentiationH. obsoletus between sympatric (including syntopic) anddelineated geographically by host distant- populationsCa-Ud Vacinto threeCf phylogenetic lineages H. obsoletusplant preference: sensu lato 1) , 2) i 3) . Cryptic species diversification within - is supported by the genetic divergence of mitochondrial genes (1.1% 1.5%), microsatellites that confirm population- clustering into three genetic groups associated with specific- host plantH. obsoletus (average assignment >90%) and genetic distances among host associated metapopulations- Hyalesthes comparable thra tocicus the most(2.1% closely - related, morphologically H.distinguishable obsoletus sensu species lato differentiation on the 3.3%). Consequences'Ca. of the epidemiology of P. solani' arein situassessed by multilocus genotyping of the phytoplasmaMediter isolates detected (1) in natural epidemiological systems of the ranean vineyards and (2) plantCa. inoculation experiments using naturally infectedtuf-a/-ab vector and tufpopulations.- Basic ' P. solani' epidemiologicalUd and Ca divergence into b cycles, each associatedVac -with tuf- , respectively, is confirmed with evidencestuf- of intermixingo induced br epidemiology, while complexity of the ofb epidemiol H.gy obsoletus is further ex evidenced C. foetida andth ough its dual influence host-plant of the . third genetic lineage the vector KEY WORDS Hyalesthes obsoletus, host- : plant, population genetics, cryptic SCIENTIFICevolution, stolbur FIELD phytoplasma,: Biology epidemiology, multilocus genotypisation SPECIFIC SCIENTIFIC FIELD: y UDC: [[577.832+595.753] Entomolog] :576.89 :579.887(043.3) Sadržaj 1. Uvod .................................................................................................................. 1 1.1 Hyalesthes obsoletus Signoret, 1865 .......... 3 1.2 Candidatus ................................... 7 Taksonomija, biologija i vektorska uloga 1.3 'Candidatus Biologija i epidemiologija fitoplazmi (' Phytoplasma') ........................................................... 11 Phytoplasma solani' − stolbur fitoplazma: epidemiologija uslovljena 1.4 H. obsoletus vektorom, putevi prenošenja i molekularna identifikacija Ca. P. solani' ..................................................................................................................... 14 Specijalizacija vektora prema biljkama domaćinima i uticaj na 2. epidemiologiju Ciljevi ................................ ' ...................................................................................... 17 3. Materijal i metode .............................................................................................. 18 3.1 - H. obsoletus...................................................... 19 3.1.1 ............................... 19 Populaciono genetičke analize diverziteta 3.1.2 ............................................................................................................................. 19 Sakupljanje populacija specijalizovanih prema biljci domaćinu 3.1.3 ............................................................................ 20 Ekstrakcija DNK 3.1.3.1 ..................................................................... 20 Genotipizacija mitohondrijskih markera 3.1.3.2 Amplifikacija ................................ mitohondrijskih................................ markera......................................................................... 23 3.1.3.3 FilogenetskeGenealogija analizemitohondrijskih ................................ gena,................................ analize diverziteta................................ i populaciona........... 27 diferencijacija 3.1.4 .................................................... 28 3.1.4.1 ............................................
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