ࡱ>    bjbjqq7eeW 8PLiV, "fShShShShShShS$X7[SU$$$SU@P<P<P<$bfSP<$fSP<P<FFLbl <9KRS!VHiVK|/\<9/\ FLFL0/\vLbP<'!C"SS;XiV$$$$/\  : Oligomer RNA, sposb regulowania procesu powstawania miRNA oraz zastosowanie oligomerw RNA jako regulatorw procesu powstawania miRNA Przedmiotem rozwizania s oligomer RNA, sposb regulowania procesu powstawania miRNA oraz zastosowanie oligomerw RNA jako regulatorw procesu powstawania miRNA. Bardziej szczegBowo, rozwizanie dotyczy zastosowania oligomerw RNA zaburzajcych struktur pre-miRNA jako regulatorw procesu powstawania miRNA. OddziaBywania oligomeru z prekursorem miRNA (pre-miRNA) zmieniaj struktur drugo- i trzeciorzdow prekursora. W rezultacie, pre-miRNA nie jest specyficznie rozpoznawany i city przez rybonukleaz Dicer i w konsekwencji nie dochodzi do powstawania okre[lonego miRNA. Czsteczki miRNA peBni bardzo istotne role zarwno w procesach fizjologicznych (np. procesach: rozwojowych, r|nicowania komrek, apoptozy, utrzymywania linii komrek macierzystych czy morfogenezie mzgu) jak i patologicznych (transformacja nowotworowa, procesy neurodegeneracyjne itp.)  ADDIN EN.CITE Mattick2005590591580926414 Spec No 12005Apr 15Small regulatory RNAs in mammalsR121-32ARC Special Research Centre for Functional and Applied Genomics, Institute for Molecular Bioscience, University of Queensland, Brisbane, Australia. j.mattick@imb.uq.edu.auMattick, J. S.Makunin, I. V.Hum Mol GenetAnimalsChromatin/metabolism*Gene Expression RegulationHumansInsulin/metabolismMammalsMicroRNAs/*geneticsModels, BiologicalPhenotypeRNA/metabolismRNA Editing/geneticsRNA SplicingRNA, Double-Stranded/*geneticsRNA, Small Interfering/*geneticsRNA, Small Nucleolar/*geneticsTime FactorsTissue DistributionTranscription, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=15809264Croce200511301131600912612212005Jul 15miRNAs, cancer, and stem cell division6-7Department of Molecular Virology, Immunology, and Medical Genetics and Comprehensive Cancer Center, The Ohio State University, Columbus, Ohio 43210, USA.Croce, C. M.Calin, G. A.CellAnimalsDisease ProgressionGene Expression ProfilingGene Expression RegulationHumans*MicroRNAs/genetics/metabolism*Neoplasms/genetics/metabolism*Stem Cells/cytology/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=16009126Giraldez200511601161577472230857232005May 6MicroRNAs regulate brain morphogenesis in zebrafish833-8Developmental Genetics Program, Skirball Institute of Biomolecular Medicine and Department of Cell Biology, New York University School of Medicine, New York, NY 10016, USA. giraldez@saturn.med.nyu.eduGiraldez, A. J.Cinalli, R. M.Glasner, M. E.Enright, A. J.Thomson, J. M.Baskerville, S.Hammond, S. M.Bartel, D. P.Schier, A. F.ScienceAnimalsBody PatterningBrain/*embryologyCell DifferentiationCentral Nervous System/embryologyGastrula/physiologyGene SilencingHeart/embryologyMicroRNAs/genetics/metabolism/*physiology*MorphogenesisMutationNeurons/cytologyPhenotypeRNA Processing, Post-TranscriptionalRNA, Double-Stranded/metabolismRibonuclease III/genetics/metabolismSignal TransductionSomites/cytology/physiologySpinal Cord/embryologyZebrafish/*embryology/*geneticshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=15774722[1-3]. Wyniki ostatnich badaD wskazuj, i| ponad 60% ludzkich genw znajduje si pod kontrol miRNA [4]. Odczytanie informacji zapisanej w genomach organizmw |ywych to jedno z najwa|niejszych zadaD stawianych wspBczesnej biologii molekularnej i bioinformatyce. Wykazano, |e pojedynczy gen mo|e by zrdBem wicej ni| jednego transkryptu. Co wicej z pojedynczego transkryptu mo|e powsta wiele r|nych mRNA. Nie stwierdzono, by istniaBa prosta korelacja pomidzy wielko[ci genomu, a zBo|ono[ci organizmu. Jedynie u bakterii i prostych organizmw zwierzcych lub ro[linnych wikszo[ genomu koduje biaBka. U organizmw wy|szych sekwencje kodujce biaBka stanowi jedynie niewielk cz[ materiaBu genetycznego (u czBowieka mniej ni| 5%). W wyniku poznania peBnej sekwencji genomu ludzkiego stwierdzono, i| wbrew wcze[niejszym oczekiwaniom nie zawiera on 150 tysicy, a jedynie okoBo 25 tysicy genw, a wic mniej wicej tyle samo, co prosta ro[lina modelowa Arabidopsis thaliana. Jednak prawdziwy przeBom w nauce w odniesieniu do mechanizmw rzdzcych procesem ekspresji informacji genetycznej dokonaB si dopiero w ostatnich latach, gBwnie za spraw odkrycia zjawiska interferencji RNA (RNAi, ang. RNA interference) oraz krtkich regulatorowych RNA, w tym mikroRNA (miRNA)  ADDIN EN.CITE Fire199840040948665339166691998Feb 19Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans806-11Carnegie Institution of Washington, Department of Embryology, Baltimore, Maryland 21210, USA. fire@mail1.ciwemb.eduFire, A.Xu, S.Montgomery, M. K.Kostas, S. A.Driver, S. E.Mello, C. C.NatureAnimalsCaenorhabditis elegans/drug effects/*genetics*Caenorhabditis elegans ProteinsCalmodulin-Binding Proteins/geneticsGene Expression Regulation/*drug effectsGenes, HelminthHelminth Proteins/geneticsMuscle Proteins/geneticsPhenotypeRNA, Antisense/pharmacologyRNA, Double-Stranded/*pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9486653[5]. OkazaBo si, i| w genomach organizmw eukariotycznych, w tym i w genomie czBowieka, obecne s sekwencje kodujce regulatorowe czsteczki RNA tzw. miRNA. Sekwencje takie mog wystpowa w obrbie genw kodujcych biaBka (czsto w intronach), jak i tworzy niezale|ne geny nie kodujce biaBek, a jedynie RNA. Transkrypty kodujce miRNA zwane s pri-miRNA. W ich strukturze wystpuj zazwyczaj okoBo 50-80 nukleotydowe, nie w peBni komplementarne rejony dwuniciowe, ktre ze wzgldu na swj ksztaBt okre[lane s mianem struktur typu spinki do wBosw (ang. hairpin). Stwierdzono, i| miRNA powstaj zarwno w organizmach ro[linnych, jak i zwierzcych, jednak proces ich dojrzewania w obu typach komrek jest wyraznie r|ny. W przypadku komrek ludzkich wyr|niamy trzy podstawowe etapy tworzenia miRNA  ADDIN EN.CITE Bartel200411101111474443811622004Jan 23MicroRNAs: genomics, biogenesis, mechanism, and function281-97Whitehead Institute for Biomedical Research, 9 Cambridge Center, Cambridge, MA 02142, USA. dbartel@wi.mit.eduBartel, D. P.CellAnimalsBase SequenceCaenorhabditis elegansDrosophila*GenomeHumansMicroRNAs/*physiologyModels, BiologicalMolecular Sequence DataPhenotypePlant Proteins/physiologyProtein BiosynthesisRNA, Small Interfering/metabolismSpecies SpecificityTranscription, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14744438[6]. W pierwszym, enzym Drosha wycina z pri-miRNA dwuniciowy fragment przyjmujcy struktur typu spinki do wBosw. W drugim etapie nowo powstaBa 50-80 nukleotydowa czsteczka - pre-miRNA transportowana jest z jdra komrkowego do cytoplazmy przez eksportyn 5. W trzecim etapie, pre-miRNA rozpoznawany jest przez rybonukleaz Dicer, ktra wycina z niego 20-23 nukleotydowy dupleks. Nastpnie dupleks ten jest przenoszony do innego kompleksu biaBkowego  RISC (ang. RNA-induced silencing complex). W kolejnym etapie RISC zostaje zaktywowany. Proces ten polega na usuniciu jednej z nici dupleksu, podczas gdy druga, zwana miRNA, pozostaje w kompleksie i sBu|y jako sonda umo|liwiajca specyficzne rozpoznawanie w peBni lub cz[ciowo komplementarnych czsteczek RNA  ADDIN EN.CITE <EndNote>Nykanen2001550551170112210732001Nov 2ATP requirements and small interfering RNA structure in the RNA interference pathway309-21Department of Biochemistry and Molecular Pharmacology, University of Massachusetts Medical School, 55 Lake Avenue North, Worcester, MA 01655, USA.Nykanen, A.Haley, B.Zamore, P. D.CellAdenosine Triphosphate/*metabolism/physiologyAnimalsDrosophila/embryology*Gene SilencingModels, GeneticPhosphorylationProteins/metabolism*RNA Processing, Post-TranscriptionalRNA, Double-Stranded/*metabolismRNA, Small InterferingRNA, Untranslated/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11701122Hammond200021802181074921340467752000Mar 16An RNA-directed nuclease mediates post-transcriptional gene silencing in Drosophila cells293-6Genetica, Inc., Cold Spring Harbor, New York 11724, USA.Hammond, S. M.Bernstein, E.Beach, D.Hannon, G. J.NatureAnimalsAntigens, CD8/geneticsCell Cycle/geneticsCells, CulturedCyclin E/geneticsDrosophilaEndoribonucleases/*metabolism*Gene SilencingHumansLac OperonRNA, Double-Stranded/*physiologyRNA, Messenger/metabolismSubstrate SpecificityTranscription, GeneticTransfectionhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10749213</URL></MDL></Cite></EndNote>[7,8]. Aktywny kompleks RISC mo|e wiza si z mRNA. Je|eli mRNA jest w peBni komplementarny do znajdujcego si w RISC miRNA, wwczas zostaje on przecity (mniej wicej w [rodku dupleksu mRNA/miRNA)  ADDIN EN.CITE <EndNote><Cite><Author>Haley</Author><Year>2004</Year><RecNum>174</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><REFNUM>174</REFNUM><ACCESSION_NUMBER>15170178</ACCESSION_NUMBER><VOLUME>11</VOLUME>72004JulKinetic analysis of the RNAi enzyme complex599-606Department of Biochemistry and Molecular Pharmacology, University of Massachusetts Medical School, Worcester, Massachusetts 01605, USA.Haley, B.Zamore, P. D.Nat Struct Mol BiolAdenosine Triphosphate/metabolismAnimalsBase SequenceCatalysisDNA PrimersDrosophila melanogasterKineticsProtein Binding*RNA InterferenceRNA, Small Interfering/metabolismSequence Homology, Nucleic Acidhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=15170178[9]. W konsekwencji, w stosunkowo krtkim czasie caBa pula mRNA zostaje zdegradowana, ma miejsce tzw. potranskrypcyjne wyciszanie genw (PTGS; ang. posttranscriptional gene silencing). Je[li jednak mRNA jest tylko cz[ciowo komplementarny do wystpujcego w RISC miRNA wwczas nie ulega on degradacji, a pozostaje zwizany z kompleksem biaBkowym, przez co nie mo|e sBu|y jako matryca do syntezy biaBka. Dochodzi zatem do wyciszenia genu na poziomie translacji (TR ang. translational repression). Rybonukleaza Dicer nale|y do rodziny RNaz III, tj. do endorybonukleaz, ktre specyficznie tn dwuniciowe czsteczki RNA. Wszystkie enzymy nale|ce do tej rodziny charakteryzuje obecno[ jednej lub dwch domen rybonukleazowych, zwyczajowo zwanych domen RNazy III. Na skutek trawienia dsRNA przez RNazy III powstaj specyficzne produkty posiadajce grup fosforanow na koDcu 5 i dwa niesparowane nukleotydy na koDcu 3 . RNazy III r|ni si wielko[ci, maj od 200  2000 aminokwasw. Zebrane dotychczas dane sugeruj, i| w genomie czBowieka zakodowana jest tylko jedna nukleaza Dicer. W przypadku biogenezy miRNA, rybonukleaza Dicer odpowiedzialna jest za wycinanie z prekursorowych pre-miRNA krtkich okoBo 20-23 nukleotydowych dupleksw zawierajcych miRNA. W oparciu o dane biochemiczne stwierdzono, |e ludzka rybonukleaza Dicer i bakteryjna RNaza III zawieraj tylko jedno centrum aktywne, w ktrym dochodzi do cicia obu nici RNA  ADDIN EN.CITE <EndNote><Cite><Author>Zhang</Author><Year>1997</Year><RecNum>153</RecNum><MDL><REFERENCE_TYPE>0153939104394251997Dec 9Regulation of ribonuclease III processing by double-helical sequence antideterminants13437-41Department of Biological Sciences, Wayne State University, Detroit, MI 48202, USA.Zhang, K.Nicholson, A. W.Proc Natl Acad Sci U S ABase SequenceBinding SitesEndoribonucleases/*metabolismEscherichia coli/genetics/metabolism*Escherichia coli ProteinsMolecular Sequence DataMutagenesis, Site-DirectedNucleic Acid Conformation*RNA Processing, Post-TranscriptionalRNA, Bacterial/chemistry/genetics/metabolismRNA, Double-Stranded/chemistry/genetics/metabolismRNA, Transfer, Amino Acid-Specific/chemistry/genetics/metabolismRibonuclease IIISubstrate Specificityhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9391043Macrae200615601561641051731157582006Jan 13Structural basis for double-stranded RNA processing by Dicer195-8Department of Molecular and Cell Biology, University of California, Berkeley, CA 94720, USA.Macrae, I. J.Zhou, K.Li, F.Repic, A.Brooks, A. N.Cande, W. Z.Adams, P. D.Doudna, J. A.ScienceAmino Acid SequenceAnimalsConserved SequenceCrystallography, X-RayGiardia lamblia/enzymologyHumansLanthanoid Series Elements/metabolismModels, MolecularMolecular Sequence DataProtein Structure, TertiaryRNA InterferenceRNA, Double-Stranded/*metabolismRNA, Protozoan/metabolismRecombinant Fusion Proteins/genetics/metabolismRibonuclease III/*chemistry/metabolismSchizosaccharomyces/geneticsStructure-Activity Relationshiphttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=16410517[10, 11]. Wykazano jednak, |e w przypadku ludzkiej Dicer, wchodzce w jej skBad dwie domeny RNazy III dziaBaj niezale|nie (ka|da tnie tylko jedn z nici)  ADDIN EN.CITE <EndNote><Cite><Author>Macrae</Author><Year>2006</Year><RecNum>156</RecNum><MDL><REFERENCE_TYPE>01561641051731157582006Jan 13Structural basis for double-stranded RNA processing by Dicer195-8Department of Molecular and Cell Biology, University of California, Berkeley, CA 94720, USA.Macrae, I. J.Zhou, K.Li, F.Repic, A.Brooks, A. N.Cande, W. Z.Adams, P. D.Doudna, J. A.ScienceAmino Acid SequenceAnimalsConserved SequenceCrystallography, X-RayGiardia lamblia/enzymologyHumansLanthanoid Series Elements/metabolismModels, MolecularMolecular Sequence DataProtein Structure, TertiaryRNA InterferenceRNA, Double-Stranded/*metabolismRNA, Protozoan/metabolismRecombinant Fusion Proteins/genetics/metabolismRibonuclease III/*chemistry/metabolismSchizosaccharomyces/geneticsStructure-Activity Relationshiphttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=16410517</URL></MDL></Cite></EndNote>[11]. Zgodnie z komputerowymi symulacjami prekursory pre-miRNA przyjmuj struktur RNA typu spinki, ktra obejmuje: i) rejon dwuniciowy (trzon spinki), zawierajcy elementy destabilizujce w postaci ptli wewntrznych, jednostronnych wybrzuszeD i niesparowaD oraz ii) rejon jednoniciowy (ptla terminala), Bczcy sekwencje tworzce trzon spinki. Pierwsze doniesienia dotyczce istotnego znaczenia cech struktury prekursorw miRNA w procesie powstawania miRNA wskazywaBy, i| kluczow rol podczas procesu rozpoznawania pre-miRNA przez rybonukleaz Dicer peBni charakterystyczna struktura i niesparowane nukleotydy na koDcu 3 prekursora miRNA (obecno[ dwch niesparowanych nukleotydw na koDcu 3 ) [12]. Uwa|ano, i| ptla terminalna nie ma wikszego wpBywu na proces tworzenia miRNA [13]. Han i wspBautorzy pokazali jednak, i| od wielko[ci ptli terminalnej zale|y wydajno[ wizania niektrych biaBek, uczestniczcych w procesie powstawania miRNA, do prekursorw. Ponadto autorzy zaobserwowali, i| z obecno[ci wikszych ptli terminalnych w prekursorach wi|e si wiksza wydajno[ powstawania pre-miRNA oraz miRNA. Podobn zale|no[ zaobserwowali Zhang X. i Zeng Y. [14]. Na podstawie uzyskanych wynikw autorzy wysnuli hipotez, i| du|e ptle terminalne wpBywaj korzystnie na elastyczno[ struktury drugo- i trzeciorzdowej prekursorw (pri- oraz pre-miRNA), co z kolei korzystnie wpBywa na szybko[ uwalniania produktw powstajcych w wyniku cicia prekursorw miRNA przez rybonukleazy Drosha oraz Dicer (wzrasta wydajno[ produkcji miRNA). Kolejne doniesienia opublikowane przez Michlewskiego G. i wspBautorw wskazywaBy na kluczowy udziaB ptli terminalnych obecnych w prekursorach pri-miRNA w procesie regulacji powstawania miRNA [15]. Autorzy pokazali, i| zastosowanie 16-nukleotydowych oligomerw RNA, komplementarnych do sekwencji obecnych w ptlach terminalnych pri-miRNA (LooptomiRw), zapobiegaBo wizaniu si kluczowych biaBkowych czynnikw regulatorowych z tym rejonem, co w konsekwencji powodowaBo zahamowanie powstawania pre-miRNA. Ponadto, Kloosterman i wspBpracownicy pokazali, i| powstawanie wybranych miRNA mo|e zosta zahamowane poprzez podanie krtkich czsteczek RNA (oraz ich pochodnych) komplementarnych do sekwencji RNA obecnych w prekursorach miRNA, stanowicych miejsca ci dla rybonukleaz Drosha oraz Dicer [16]. W zgBoszeniu patentowym P-384455 (opubl. 2009-08-17) wykazano, i| aptamery RNA wi|ce ludzk rybonukleaz Dicer, mog inhibowa aktywno[ tego enzymu na zasadzie kompetycyjnej oraz allosterycznej [17]. Wyniki ostatnio prowadzonych badaD wskazuj, i| przedstawione poni|ej rozwizanie sposobu inhibicji procesu powstawania miRNA poprzez zastosowanie oligomerw RNA (oraz ich pochodnych) wi|cych specyficznie wybrane prekursory miRNA i zaburzajcych ich struktur, umo|liwia projektowanie i tworzenie czsteczek inhibitorowych selektywnie hamujcych powstawanie wybranych czsteczek miRNA. Obecnie znana jest sekwencja ponad 1500 r|nych ludzkich czsteczek miRNA (hsa-miRNA). Czsteczki miRNA, jak si przypuszcza, bior udziaB w kontroli ekspresji ponad 60% ludzkich genw. Wiadomo rwnie|, i| geny regulowane przez miRNA odgrywaj kluczow rol w procesach rozwojowych, w powstawaniu wyspecjalizowanych komrek, tkanek i narzdw, a tak|e w programowanej [mierci komrek. Nawet niewielkie zaburzenia w procesie biogenezy miRNA, czy w procesach odpowiedzialnych za prawidBowe dziaBanie tych czsteczek, mog uruchamia szereg niekorzystnych procesw, np. nowotworowych, degeneracyjnych czy zaburzeD rozwojowych, prowadzcych do dysfunkcji narzdw i organw, a czsto [mierci organizmu. Pomimo istniejcych rozwizaD jednym z najwikszych wyzwaD stawianych wspBczesnym naukom biomedycznym jest projektowanie i otrzymywanie terapeutykw dziaBajcych w sposb selektywny na okre[lone cele biologiczne, np. i) zwizkw niszczcych wybirczo komrki nowotworowe, a nie majcych negatywnego wpBywu na komrki zdrowe; ii) zwizkw hamujcych aktywno[ konkretnych biaBek i nie wi|cych biaBek charakteryzujcych si podobn budow i funkcj do biaBka stanowicego cel terapii. Celem wynalazku jest umo|liwienie zaprojektowania czsteczek regulatorowych, ktre zarwno w sposb uniwersalny, jak i wybirczy mog kontrolowa proces dojrzewania, a tak|e funkcjonowania wybranych (konkretnych) czsteczek miRNA. Zaproponowane rozwizanie otwiera drog do opracowania nowych, specyficznych metod terapeutycznych, ktre mog znalez zastosowanie w leczeniu schorzeD wywoBanych zmian prawidBowego poziomu ekspresji wybranych czsteczek regulatorowych, np. w leczeniu chorb nowotworowych, neurodegeneracyjnych, czy zakaznych. Przedmiotem wynalazku jest oligomer RNA charakteryzujcy si tym, |e zawiera: sekwencje komplementarne do rejonw jednoniciowych czsteczki pre-miRNA lub sekwencje komplementarne do rejonw dwuniciowych czsteczki pre-miRNA lub sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA przy czym oligomer oddziaBuje z pre-miRNA, zaburza struktur pre-miRNA i hamuje proces wycinania miRNA z pre-miRNA. Korzystnie, gdy oligomer RNA dziaBa dwuetapowo: i) jako inhibitor kompetycyjny, na etapie wizania z rybonukleaz Dicer, ii) jako czsteczka zaburzajca struktur substratu pre-miRNA, na etapie wizania z substratem pre-miRNA. Korzystnie, gdy oligomer RNA dziaBa jednoetapowo jako czsteczka zaburzajca struktur substratu pre-miRNA, na etapie wizania z substratem pre-miRNA. Korzystnie, gdy oligomer jest wybrany spo[rd: - oligomeru RNA oddziaBujcego z rybonukleaz Dicer, - oligomeru RNA oddziaBujcego z substratem pre-miRNA, - oligomeru RNA oddziaBujcego z jednoniciowymi rejonami substratu pre-miRNA, - oligomeru RNA oddziaBujcego z dwuniciowymi rejonami substratu pre-miRNA, - oligomeru RNA oddziaBujcego z jednoniciowymi oraz dwuniciowymi rejonami substratu pre-miRNA, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA w taki sposb, i| substrat nie jest specyficznie rozpoznawany przez rybonukleaz Dicer, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA w taki sposb, i| nie dochodzi do powstawania specyficznych produktw miRNA, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA w taki sposb, i| nie dochodzi do powstawania funkcjonalnych miRNA, - oligomeru RNA komplementarnego do jednoniciowych rejonw prekursora pre-miRNA, - oligomeru RNA komplementarnego do dwuniciowych rejonw prekursora pre-miRNA, - oligomeru RNA komplementarnego do jednoniciowych oraz dwuniciowych rejonw prekursora pre-miRNA, - oligomeru RNA zawierajcy modyfikacje zasad, cukru, reszt fosforanowych, przy czym oligomery s dBugo[ci do 75 nt. Korzystnie, gdy oligomer zawiera sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA, wybrane spo[rd sekwencji: seq. nr 2, seq. nr 3, seq. nr 4, seq. nr 8, seq. nr 9, seq.10, seq. nr 11. Kolejnym przedmiotem wedBug wynalazku jest sposb regulowania procesu powstawania miRNA, charakteryzujcy si tym, |e obejmuje etapy, w ktrych oligomer okre[lony powy|ej oddziaBuje z pre-miRNA, zaburza struktur pre-miRNA i hamuje proces wycinania miRNA z pre-miRNA przez ludzk rybonukleaz Dicer. Korzystnie, gdy sposb stanowi wybirczy i selektywny sposb inhibicji powstawania miRNA. Korzystnie, gdy oligomer dziaBa dwuetapowo: a) wi|e si z Dicer, b) produkt cicia oligomeru przez ludzk rybonukleaz Dicer wi|e si z fragmentem ptli terminalnej pre-miRNA i zmienia struktur drugo- i trzeciorzdow pre-miRNA, tak |e pre-miRNA nie jest specyficznie rozpoznawany i city przez ludzk rybonukleaz Dicer i nie dochodzi do powstawania miRNA. Korzystnie, gdy oligomer dziaBa jednoetapowo: tworzy kompleks z pre-miRNA i zmienia jego struktur, tak |e pre-miRNA nie jest specyficznie rozpoznawany i city przez ludzk rybonukleaz Dicer i nie dochodzi do powstawania miRNA. Korzystnie, gdy sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA, wybiera si spo[rd sekwencji: seq. nr 2, seq. nr 3, seq. nr 4, seq. nr 8, seq. nr 9, seq.10, seq. nr 11. Kolejnym przedmiotem wedBug wynalazku jest zastosowanie oligomerw RNA okre[lonych powy|ej zaburzajcych struktur pre-miRNA jako regulatorw procesu powstawania miRNA. Korzystnie, gdy sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA, wybrane s spo[rd sekwencji: seq. nr 2, seq. nr 3, seq. nr 4, seq. nr 8, seq. nr 9, seq.10, seq. nr 11. W celu lepszego zrozumienia omawianych zagadnieD rozwizania zostaBy przedstawione na rysunku, gdzie: Figura 1 przedstawia wpByw oligomeru ATD_15.52 oraz produktw cicia ATD_15.52 przez rybonukleaz Dicer na proces powstawania hsa-miR-210 Reakcje cicia prekursora pre-hsa-miR-210 przeprowadzono w obecno[ci oligorybonukleotydu ATD_15.52 oraz syntetycznie otrzymanych produktw cicia tego oligomeru przez ludzk rybonukleaz Dicer: 21-nukleotydowego fragmentu 5 (5 _ATD_15.52) oraz 35-nukleotydowego fragmentu 3 (3 _ATD_15.52). (A) W reakcjach oznaczonych numerem 1 stosunek molowy oligomer : Dicer wynosiB 1:1, w reakcjach oznaczonych numerem 2 10:1, a w reakcjach oznaczonych numerem 3 100:1. Dodatkowo zawsze przeprowadzano dwie reakcje kontrolne: (K+) - standardowa reakcja bez dodatku oligomeru oraz (K-) - standardowa reakcja bez enzymu. (B) Graficzne przedstawienie wydajno[ci cicia prekursora pre-hsa-miR-210 przez Dicer w obecno[ci czsteczek inhibitorowych. Podane warto[ci dotyczce inhibicji powstawania miRNA-210 zostaBy wyznaczone w odniesieniu do reakcji kontrolnej, przeprowadzonej bez dodatku czsteczek inhibitorowych. W przypadku reakcji kontrolnej wydajno[ cicia pre-miRNA-210 przez ludzk rybonukleaz Dicer zostaBa ustalona w oparciu o pomiar intensywno[ci sygnaBu pochodzcego od substratu - prekursora pre-miRNA oraz produktu  miRNA. Figura 2 przedstawia: (A) Autoradiogram prezentujcy wynik elektroforezy w niedenaturujcym |elu poliakrylamidowym: wyznakowanego radioizotopowo oligomeru ATD_15.52 ([cie|ka 1) oraz kompleksu ATD_15.52 : pre-hsa-miR-210 ([cie|ka 2); wyznakowanego radioizotopowo 21-nukleotydowego fragmentu 5 oligomeru ATD_15.52 ([cie|ka 3) oraz kompleksu 5 _ATD_15.52 : pre-hsa-miR-210 ([cie|ka 4); wyznakowanego radioizotopowo 35-nukleotydowego fragmentu 3 oligomeru ATD_15.52 ([cie|ka 5) oraz reakcji hybrydyzacji czsteczki 3 _ATD_15.52 oraz prekursora pre-hsa-miR-210 ([cie|ka 6). (B) Autoradiogram prezentujcy wynik pomiaru stopnia zwizania wyznakowanego radioizotopowo na koDcu 5 oligomeru ATD_15.52 z rybonukleaz Dicer. Mieszaniny reakcyjne zawierajce Dicer oraz wyznakowan radioizotopowo czsteczk ATD_15.52 nanoszone byBy na membran nitrocelulozow po: 0, 1, 2, 3, 4 i 5h inkubacji z enzymem w temperaturze 37C (w trakcie inkubacji z Dicer ATD_15.52 byB city na 21-nukleotydowy fragment 5 oraz 35-nukleotydowy fragment 3 ). Figura 3 przedstawia zaproponowane przez modelowanie komputerowe struktury analizowanych czsteczek pre-hsa-miR-210 (A) oraz kompleksu utworzonego przez prekursor pre-hsa-miR-210 oraz 21 nukleotydowy fragment 5 oligomeru ATD_15.52 (B), przyjmujce najni|sz warto[ energii swobodnej. Figura 4 przestawia analiz wpBywu 12-nukleotydowych oligomerw RNA wi|cych specyficznie rejony jednoniciowe prekursorw pre-miRNA na proces powstawania miRNA w obecno[ci ludzkiej rybonukleazy Dicer: (A) WpByw anti_Loop-16-1 na cicie pre-hsa-miR-16-1 (B) WpByw anti_Loop-33a na cicie pre-hsa-miR-33a (C) WpByw anti_Loop-210 na cicie pre-hsa-miR-210 Opis rysunku zgodny z opisem dla Figury 1(A) oraz (B). Figura 5 przedstawia wpByw oligomeru ATD_13.6 oddziaBujcego specyficznie z prekursorem pre-hsa-miR-210 na proces powstawania miRNA-210 w obecno[ci ludzkiej rybonukleazy Dicer: (A) Autoradiogram przedstawiajcy wynik elektroforezy w niedenaturujcym |elu poliakrylamidowym wyznakowanego radioizotopowo oligomeru ATD_13.6 ([cie|ka 1) oraz kompleksu ATD_13.6 : pre-hsa-miR-210 ([cie|ka 2) (B) Zaproponowane przez modelowanie komputerowe struktury analizowanych czsteczek: pre-hsa-miR-210, ATD_13.6 oraz kompleksu ATD_13.6 : pre-hsa-miR-210, przyjmujce najni|sz warto[ energii swobodnej. (C) Analiza wpBywu ATD_13.6 na cicie pre-hsa-miR-210. Opis rysunku zgodny z opisem dla Figury 1(A) oraz (B). Figura 6 przedstawia wpByw czsteczek RNA stanowicych fragmenty mRNA wi|cych specyficznie rejony jedno- i dwuniciowe prekursora pre-hsa-miR-210 na proces powstawania miRNA-210 w obecno[ci ludzkiej rybonukleazy Dicer: (A) Analiza wpBywu PCDH21_fr na cicie pre-hsa-miR-210. Czsteczka PCDH21_fr jest fragmentem mRNA kodujcego biaBko protokadheryn (zale|ne od wapnia biaBko uczestniczce w adhezji komrek). Czsteczka ta jest komplementarna do rejonu dwuniciowego prekursora pre-hsa-miR-210. (B) Analiza wpBywu THAP4_fr na cicie pre-hsa-miR-210. Czsteczka THAP4_fr jest fragmentem mRNA kodujcego biaBko THAP4 (biaBko wi|ce DNA oraz jony metali). Czsteczka ta jest komplementarna do rejonu jednoniciowego ptli terminalnej prekursora pre-hsa-miR-210. Opis rysunku zgodny z opisem dla Figury 1(A) oraz (B). Poni|ej zaprezentowano przykBadowe realizacje zdefiniowanego powy|ej wynalazku. PrzykBad Jednym z kluczowych enzymw uczestniczcych w biogenezie miRNA, jak i innych krtkich regulatorowych RNA, jest rybonukleaza Dicer. Wycina ona funkcjonalne czsteczki miRNA z dwuniciowych prekursorw pre-miRNA. Postawiono zatem pytanie czy odpowiednio zaprojektowane oligorybonukleotydy, oddziaBujce z prekursorami pre-miRNA i zaburzajce ich struktur, mog wpBywa na proces powstawania miRNA (czy proces powstawania krtkich regulatorowych RNA mo|e by kontrolowany przez inne czsteczki RNA)? Aby rozwiza ten problem testowano wpByw czsteczek RNA (oraz ich pochodnych), wi|cych specyficznie jedno- i dwuniciowe rejony prekursorw pre-miRNA na proces tworzenia odpowiednich czsteczek miRNA, w ktrym bierze udziaB ludzka rybonukleaza Dicer. Badanie wpBywu krtkich czsteczek RNA wyzwalanych przez Dicer i oddziaBujcych z pre-miRNA na proces powstawania miRNA Wcze[niejsze badania, bdce przedmiotem zgBoszenia patentowego P-0384455 (opubl. 2009-08-17), pozwoliBy ustali, i| oligomer ATD_15.52 wyraznie hamuje cicie pre-hsa-miR-210 (gdy zastosowano 100-krotny nadmiar oligomeru wobec Dicer tylko 15% substratu ulegaBo ciciu  w porwnaniu z reakcj kontroln przeprowadzon bez dodatku czsteczki inhibitorowej) [17]. Nastpnie wykazano, i| w wyniku cicia ATD_15.52 przez Dicer powstaj dwa produkty: 21 nukleotydowy fragment 5 oraz 35 nukleotydowy fragment 3 [17]. 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hZB*CJOJPJQJ^JaJnHphtH JV.xzɮx]B'5h h?B*CJOJPJQJ^JaJnHphtH5h h)B*CJOJPJQJ^JaJnHphtH5h h(SB*CJOJPJQJ^JaJnHphtH5h h-5B*CJOJPJQJ^JaJnHphtH5h h5B*CJOJPJQJ^JaJnHphtH5h hB*CJOJPJQJ^JaJnHphtH5h h".OB*CJOJPJQJ^JaJnHphtH5h h*B*CJOJPJQJ^JaJnHphtH x:::6;ɮɓ|eN757NU,h h5CJOJPJQJ^JaJnHtH,h hCJOJPJQJ^JaJnHtH,h h}@CJOJPJQJ^JaJnHtH,h h?CJOJPJQJ^JaJnHtH5h h?B*CJOJPJQJ^JaJnHphtH5h hoB*CJOJPJQJ^JaJnHphtH5h hB*CJOJPJQJ^JaJnHphtH5h h( B*CJOJPJQJ^JaJnHphtH admiar fragmentu 5 wobec Dicer, zaledwie 9% substratu ulegaBo ciciu), natomiast obecno[ fragmentu 3 praktycznie nie wywiera wpBywu na cicie prekursora pre-hsa-miR-210 (Figura 1). Kolejne badania pokazaBy, i| fragment 5 , powstajcy w wyniku przecicia czsteczki ATD_15.52 przez rybonukleaz Dicer, nie pozostaje zwizany z enzymem (Figura 2), natomiast mo|e on oddziaBywa z prekursorem pre-hsa-miR-210 (Figura 2 oraz 3). WpByw oligomeru ATD_15.52 oraz syntetycznie otrzymanych produktw cicia ATD_15.52 przez ludzk rybonukleaz Dicer: 21-nukleotydowego fragmentu 5 (5 _ATD_15.52) oraz 35-nukleotydowego fragmentu 3 (3 _ATD_15.52), na proces powstawania miRNA-210, testowano w warunkach in vitro. W tym celu przeprowadzano standardowe reakcje trawienia ludzkich pre-hsa-miR-210 przez Dicer. W pojedynczym eksperymencie, wykonanym co najmniej trzykrotnie dla ka|dego oligomeru, przeprowadzano 5 reakcji: dwie kontrolne (bez dodatku oligomeru (K+) oraz bez enzymu (K-)) oraz trzy reakcje z dodatkiem r|nych ilo[ci ATD_15.52, 5 _ATD_15.52 lub 3 _ATD_15.52 (w reakcjach tych stosunek molowy Dicer : oligomer wynosiB 1:1, 1:10 i 1:100). Sekwencje wybranych czsteczek: pre-hsa-miR-210 5'gccccugcccaccgcacacugcgcugccccagacccacugugcgugugacagcggcug3 ATD_15.52 5 gggagaaucauaaguagcgcagugagucguugugcugcccauguuaacaguuagcc3 5 _ATD_15.52 5 gggagaaucauaaguagcgca3 3 _ATD_15.52 5 gugagucguugugcugcccauguuaacaguuagcc3 Reakcje prowadzono w objto[ci 10 l w standardowym buforze dla rybonukleazy Dicer (250 mM NaCl, 20 mM Tris-HCl pH 7.5, 2.5 mM MgCl2). Do reakcji wykorzystywano pre-miRNA wyznakowane na koDcu 5 radioaktywnym fosforem. Czsteczki RNA znakowano radioizotopowo zgodnie z nastpujc procedur: - jednoniciowy RNA (ssRNA) denaturowano przez ogrzanie do 95C przez 2 min. igwaBtowne schBodzenie na lodzie (10 min). Reakcj prowadzono w objto[ci 30l wmieszaninie o skBadzie podanym w Tabeli 1. Tabela  SEQ Tabela \* ARABIC 1. SkBad mieszaniny do reakcji radioizotopowego znakowania ssRNA SkBadnikst|enie pocztkowest|enie koDcowebufor do kinazowania110x1x ATP[ 32P] 4000-5000 Ci/mmol50 Ci ssRNA100 pmoli/l0,3 pM kinazy polinukleotydowej T4 10 U/l0,5 U H2OuzupeBniano do objto[ci 30 l1SkBad buforu do kinazowania 1x: 70 mM Tris-HCl ph 7.6, 10 mM MgCl2, 5 mM DTT. - mieszanin inkubowano 30 min. w temp. 37C, - mieszanin reakcyjn rozcieDczano do objto[ci 200 l i dodawano 100 l fenolu oraz 100 l chloroformu, - wytrzsano i wirowano 1 min. z prdko[ci 14000 rpm, - zawierajc kwas nukleinowy faz wodn przenoszono do nowej probwki i dodawano 200 l chloroformu, wytrzsano i wirowano 1 min. z prdko[ci 14000 rpm (dwukrotnie), - do otrzymanej po ekstrakcji fazy wodnej dodawano 20 l 3M CH3COONa i 3 obj. 96% etanolu, - roztwr inkubowano przez noc w temperaturze (-20oC), - wirowano 20 min. z prdko[ci 14000 rpm, - roztwr dekantowano, a osad przemywano 500 l 70% etanolu, - wirowano 20 min. z prdko[ci 14000 rpm, - roztwr dekantowano, a osad suszono i rozpuszczano w 20 l H2O, - przy u|yciu aparatu Beckman okre[lano ilo[ materiaBu radioaktywnego wprowadzonego do wyj[ciowej puli RNA. Wyznakowane radioizotopowo substraty pre-miRNA renaturowano poprzez podgrzanie do 95oC a nastpnie powolne schBodzenie do temperatury pokojowej. W kolejnym etapie przeprowadzano 10 minutow wstpn inkubacj r|nej ilo[ci oligomerw z Dicer (w stosunku molowym Dicer : oligomer 1:1, 1:10 i 1:100) w buforze reakcyjnym, w temperaturze 25oC. Nastpnie dodawano pre-miRNA. Reakcj trawienia prowadzono 30 minut w temperaturze 37(C (Tabela 2). Dla ka|dej czsteczki wykonywano po trzy powtrzenia ka|dej reakcji. Tabela  SEQ Tabela \* ARABIC 2. Badanie wpBywu oligomerw na proces powstawania miRNA w obecno[ci rybonukleazy Dicer - skBad mieszaniny reakcyjnej kontrola (+)kontrola (-)123oligomer--1 l (1 pmol/l)1 l (10 pmol/l)1 l (100 pmol/l)Dicer1U-1U1U1Upre-miRNA10000 cpm10000 cpm10000 cpm10000 cpm10000 cpmBufor2 5x2 l2 l2 l2 l2 lH20do 10 ldo 10 ldo 10 ldo 10 ldo 10 l2 skBad buforu (1x): 250 mM NaCl, 20 mM Tris-HCl pH 7.5, 2.5 mM MgCl2 Po zakoDczeniu reakcji uzyskane produkty denaturowano poprzez podgrzanie do 95(C przez 5 minut i gwaBtownie schBadzano na lodzie, a nastpnie analizowano metod elektroforezy w15% denaturujcym |elu poliakrylamidowym. RozdziaB elektroforetyczny prowadzono wnastpujcych warunkach: - preelektroforeza: 1200V, 50W, 10mA przez 30 min. - elektroforeza wBa[ciwa: 1200V, 50W, 40mA przez 4h. Po rozdziale elektroforetycznym skBad mieszaniny analizowano wykorzystujc skaner materiaBw znakowanych radioizotopowo (Fuji, FLA-5100). B) Badanie wpBywu oligomerw RNA wi|cych rejony jedno- i dwuniciowe obecne w pre-miRNA na proces powstawania miRNA Biorc pod uwag wyniki opisane w przykBadzie A, wskazujce, i| oligomery specyficznie oddziaBujce z prekursorem pre-miRNA mog wydajnie hamowa proces powstawania wybranego miRNA (Figury 1, 2 oraz 3), przeprowadzono szereg reakcji trawienia prekursorw pre-miRNA przez rybonukleaz Dicer, w obecno[ci oligomerw RNA komplementarnych do rejonw jednoniciowych (w tym: ptli terminalnych, ptli wewntrznych oraz wewntrznych wybrzuszeD) oraz rejonw dwuniciowych (rejon trzonu) wybranych prekursorw. Badano wpByw 12-nukleotydowych oligomerw RNA, specyficznie wi|cych rejony jednoniciowe prekursorw: pre-hsa-miR-16-1, -33a oraz -210, na proces powstawania odpowiednich miRNA (Figura 4). Ponadto, analizowano wpByw 56-nukleotydowego oligomeru ATD_13.6, oddziaBujcego specyficznie z prekursorem pre-hsa-miR-210 na proces powstawania miRNA-210 w obecno[ci ludzkiej rybonukleazy Dicer (Figura 5). Zgodnie z danymi umieszczonymi w publikacji [17] oligomer ATD_13.6 wi|e si rwnie| z ludzk rybonukleaz Dicer, ale nie jest city przez ten enzym. Przeprowadzono rwnie| reakcje trawienia pre-hsa-miR-210 przez Dicer w obecno[ci czsteczek THAP4_fr oraz PCDH21_fr, (stanowicych fragmenty wybranych czsteczek mRNA), wi|cych specyficznie odpowiednio rejony jedno- i dwuniciowe prekursora pre-hsa-miR-210 (Figura 6). WpByw oligomerw RNA komplementarnych do rejonw jedno- oraz dwuniciowych prekursorw pre-miRNA na proces powstawania wybranych miRNA testowano w warunkach in vitro. W tym celu przeprowadzano standardowe reakcje trawienia ludzkich pre-miRNA przez rybonukleaz Dicer. W pojedynczym eksperymencie, wykonanym co najmniej trzykrotnie dla ka|dego oligomeru, przeprowadzano 5 reakcji: dwie kontrolne (bez dodatku oligomeru oraz bez enzymu) oraz trzy reakcje z dodatkiem r|nych ilo[ci oligomeru (w reakcjach tych stosunek molowy pre-miRNA : oligomer wynosiB 1:1, 1:10 i 1:100). Sekwencje wybranych czsteczek: pre-hsa-miR-16-1 5 uagcagcacguaaauauuggcguuaagauucuaaaauuaucuccaguauuaacugugcugcugaa3 pre-hsa-miR-33a 5'gugcauuguaguugcauugcauguucuggugguacccaugcaauguuuccacagugcauca3 pre-hsa-miR-210 5'gccccugcccaccgcacacugcgcugccccagacccacugugcgugugacagcggcug3 anti_Loop-16-1 5 gaaucuuaacgc3 anti_Loop-33a 5 ggguaccaccag3 anti_Loop-210 5 ggggcagcgcag3 ATD_13.6 5 gggagaaucauaaguagcggugugugagucguggugccccauguuaacaguuagcc3 THAP4_fr 5 ggagaagcaguaggagugcagugagucgaugagcu3 PCDH21_fr 5 ggaagaacagaaguagaggugugugaguc3 Reakcje prowadzono w objto[ci 10 l, w komercyjnym buforze dla rybonukleazy Dicer. Do reakcji wykorzystywano pre-miRNA wyznakowane na koDcu 5 radioaktywnym fosforem (znakowanie przeprowadzano zgodnie z wcze[niej opisan procedur). Wyznakowane radioizotopowo substraty pre-miRNA renaturowano poprzez podgrzanie do 95oC a nastpnie powolne schBodzenie do temperatury pokojowej. W kolejnym etapie przeprowadzano 10 minutow wstpn inkubacj r|nej ilo[ci oligomerw z Dicer (w stosunku molowym Dicer : oligomer 1:1, 1:10 i 1:100) w buforze reakcyjnym, w temperaturze 25oC. Nastpnie dodawano pre-miRNA. Reakcj trawienia prowadzono 30 minut w temperaturze 37(C (Tabela 2). Dla ka|dego oligomeru wykonywano po trzy powtrzenia ka|dej reakcji. Po zakoDczeniu reakcji uzyskane produkty denaturowano poprzez podgrzanie do 95(C przez 5 minut i gwaBtownie schBadzano na lodzie, a nastpnie analizowano metod elektroforezy w15% denaturujcym |elu poliakrylamidowym. RozdziaB elektroforetyczny prowadzono wnastpujcych warunkach: - preelektroforeza: 1200V, 50W, 10mA przez 30 min. - elektroforeza wBa[ciwa: 1200V, 50W, 40mA przez 4h. Po rozdziale elektroforetycznym skBad mieszaniny analizowano wykorzystujc skaner materiaBw znakowanych radioizotopowo (Fuji, FLA-5100). Poni|ej przedstawiono wyniki omwionych powy|ej do[wiadczeD. Tabela 3. Hamowanie cicia wybranych substratw pre-miRNA przez ludzk rybonukleaz Dicer w obecno[ci oligomerw RNA substratinhibitor (oligomer)stosunek molowy enzym:inhibitorstopieD inhibicji cicia substratu [%] pre-hsa-miR-210 ATD_13.61:113%1:1054%1:10089% ATD_15.521:120%1:1052%1:10085% 5 _ATD_15.521:136%1:1079%1:10091% 3 _ATD_15.521:14%1:109%1:10015% 5 +3 _ATD_15.521:115%1:1063%1:10089% PCDH21_fr1:115%1:1040%1:10060% THAP4_fr1:113%1:1025%1:10056% anti_Loop-2101:178%1:1083%1:10094% pre-hsa-miR-16-1 anti_Loop-16-11:151%1:1083%1:10091% pre-hsa-miR-33a anti_Loop-33a1:172%1:1091%1:10098%Uzyskane wyniki umo|liwiaj stwierdzenie, i| badane oligomery RNA hamuj powstawanie miRNA wedBug dwch r|nych mechanizmw. W pierwszym przypadku, czsteczka inhibitorowa stanowi substrat dla ludzkiej rybonukleazy Dicer, a produkty wyzwalane przez Dicer hamuj powstawanie miRNA (przypadek opisany w pkt. A), natomiast w drugim przypadku, czsteczka inhibitorowa mo|e, ale nie musi, oddziaBywa z Dicer  przy czym je[li oddziaBuje z Dicer, to nie jest cita przez t rybonukleaz (przypadek opisany w pkt. B; zgodnie z danymi umieszczonymi w publikacji [17], dotyczcymi oligomeru ATD_13.6, oligomer ten wi|e si z ludzk rybonukleaz Dicer, ale nie jest city przez ten enzym). W obu przypadkach wymogiem koniecznym jest oddziaBywanie oligomeru (lub jego fragmentw) z prekursorem miRNA. W wyniku przeprowadzonych badaD ustalono, i| oligomer ATD_15.52 dziaBa dwuetapowo. W pierwszym etapie oligomer ATD_15.52 wi|e si z Dicer efektywniej od substratu (pre-hsa-miR-210), std to on jest trawiony, a nie pre-miRNA. W drugim etapie, produkt cicia oligomeru ATD_15.52 przez ludzk rybonukleaz Dicer (fragment 5 ATD_15.52), wi|e si z fragmentem ptli terminalnej prekursora pre-hsa-miR-210 zmieniajc struktur drugo- i trzeciorzdow prekursora. W konsekwencji, prekursor pre-miRNA nie jest specyficznie rozpoznawany i city przez rybonukleaz Dicer. Oligomer ATD_15.52 jest zatem inhibitorem dwufunkcyjnym. Z jednej strony, wi|c si z enzymem dziaBa jako inhibitor kompetycyjny. Z drugiej strony, wi|c si z substratem mo|e zmienia jego struktur w taki sposb, i| substrat nie mo|e zosta przyBczony do centrum aktywnego enzymu. Rezultatem podania oligomeru ATD_15.52 do reakcji trawienia prekursora pre-hsa-miR-210 przez Dicer jest zahamowanie procesu powstawania miRNA-210. Dziki specyficznym oddziaBywaniom pomidzy oligomerem i wybranym prekursorem (komplementarno[ sekwencji), proces inhibicji jest wybirczy i dotyczy selektywnego hamowania powstawania danego miRNA (Figury 1-3). Po drugie, testowane oligomery RNA, zawierajce sekwencje komplementarne do jedno- oraz dwuniciowych rejonw wybranych prekursorw miRNA, w wyniku oddziaBywania z prekursorami (tworzenie kompleksu z prekursorem), zmieniaj ich struktur w taki sposb, i| substraty nie mog by w dalszym cigu specyficznie rozpoznawane i cite przez rybonukleaz Dicer i nie dochodzi do powstawania wybranych miRNA (Figury 4-6). W zastosowanych warunkach reakcyjnych wydajna inhibicja powstawania wybranych czsteczek miRNA obserwowana byBa nawet przy najni|szym st|eniu oligomeru RNA (1 pmol oligomeru RNA, st|enie koDcowe w mieszaninie reakcyjnej  100 nM). Ponadto, zaobserwowano, |e wraz ze wzrostem st|enia oligomeru wydajno[ procesu inhibicji wzrasta. Zaprezentowane rozwizanie sposobu inhibicji procesu powstawania miRNA poprzez zastosowanie oligomerw RNA (oraz ich pochodnych) wi|cych specyficznie wybrane prekursory miRNA, umo|liwia projektowanie i tworzenie czsteczek inhibitorowych selektywnie hamujcych powstawanie wybranych czsteczek miRNA. ZgBaszajcy: Instytut Chemii Bioorganicznej PAN PeBnomocnik: dr Aleksandra Twardowska rzecznik patentowy Lista sekwencji: Sekwencja nr 1 pre-hsa-miR-210 5'gccccugcccaccgcacacugcgcugccccagacccacugugcgugugacagcggcug3 Sekwencja nr 2 ATD_15.52 5 gggagaaucauaaguagcgcagugagucguugugcugcccauguuaacaguuagcc3 Sekwencja nr 3 5 _ATD_15.52 5 gggagaaucauaaguagcgca3 Sekwencja nr 4 3 _ATD_15.52 5 gugagucguugugcugcccauguuaacaguuagcc3 Sekwencja nr 5 pre-hsa-miR-16-1 5 uagcagcacguaaauauuggcguuaagauucuaaaauuaucuccaguauuaacugugcugcugaa3 Sekwencja nr 6 pre-hsa-miR-33a 5'gugcauuguaguugcauugcauguucuggugguacccaugcaauguuuccacagugcauca3 Sekwencja nr 7 pre-hsa-miR-210 5'gccccugcccaccgcacacugcgcugccccagacccacugugcgugugacagcggcug3 Sekwencja nr 8 anti_Loop-16-1 5 gaaucuuaacgc3 Sekwencja nr 9 anti_Loop-33a 5 ggguaccaccag3 Sekwencja nr 10 anti_Loop-210 5 ggggcagcgcag3 Sekwencja nr 11 ATD_13.6 5 gggagaaucauaaguagcggugugugagucguggugccccauguuaacaguuagcc3 Sekwencja nr 12 THAP4_fr 5 ggagaagcaguaggagugcagugagucgaugagcu3 Sekwencja nr 13 PCDH21_fr 5 ggaagaacagaaguagaggugugugaguc3 Literatura:  ADDIN EN.REFLIST  1. Mattick, J.S. and I.V. Makunin, Small regulatory RNAs in mammals. Hum Mol Genet, 2005. 14 Spec No 1: p. R121-32. 2. Croce, C.M. and G.A. Calin, miRNAs, cancer, and stem cell division. Cell, 2005. 122(1): p. 6-7. 3. Giraldez, A.J., et al., MicroRNAs regulate brain morphogenesis in zebrafish. Science, 2005. 308(5723): p. 833-8. 4. Olejniczak, M., et al., Sequence-non-specific effects of RNA interference triggers and microRNA regulators. Nucleic Acids Res., 2010. 38(1): p. 1-16. 5. Fire, A., et al., Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans. Nature, 1998. 391(6669): p. 806-11. 6. Bartel, D.P., MicroRNAs: genomics, biogenesis, mechanism, and function. Cell, 2004. 116(2): p. 281-97. 7. Nykanen, A., B. Haley, and P.D. Zamore, ATP requirements and small interfering RNA structure in the RNA interference pathway. Cell, 2001. 107(3): p. 309-21. 8. Hammond, S.M., et al., An RNA-directed nuclease mediates post-transcriptional gene silencing in Drosophila cells. Nature, 2000. 404(6775): p. 293-6. 9. Haley, B. and P.D. Zamore, Kinetic analysis of the RNAi enzyme complex. Nat Struct Mol Biol, 2004. 11(7): p. 599-606. 10. Zhang, K. and A.W. Nicholson, Regulation of ribonuclease III processing by double-helical sequence antideterminants. Proc Natl Acad Sci U S A, 1997. 94(25): p. 13437-41. 11. Macrae, I.J., et al., Structural basis for double-stranded RNA processing by Dicer. Science, 2006. 311(5758): p. 195-8. 12. Zeng,Y. and B.R. Cullen, Sequence requirements for microRNA processing and function in human cells. RNA, 2003. 9(1): p. 112 23. 13. Han, J., et al., Molecular Basis for the Recognition of Primary microRNAs by the Drosha-DGCR8 Complex. Cell, 2006. 125(5): p. 887-901. 14. Zhang, X. and Y. Zeng Y, The terminal loop region controls microRNA processing by Drosha and Dicer. Nucleic Acids Res., 2010. 38(21): p. 7689-97. 15. Michlewski, G., et al., Posttranscriptional regulation of miRNAs harboring conserved terminal loops. Mol Cell., 2008. 32(3): p. 383-93. 16. Kloosterman, W.P., et al., Targeted inhibition of miRNA maturation with morpholinos reveals a role for miR-375 in pancreatic islet development. PLoS Biol., 2007. 5(8): p. e203. 17. Tyczewska, A., et al., Selection of RNA oligonucleotides that can modulate human Dicer activity in vitro. Nucleic Acid Ther., 2011. Praca w druku.  Zastrze|enia patentowe 1. Oligomer RNA znamienny tym, |e zawiera: sekwencje komplementarne do rejonw jednoniciowych czsteczki pre-miRNA lub sekwencje komplementarne do rejonw dwuniciowych czsteczki pre-miRNA lub sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA przy czym oligomer oddziaBuje z pre-miRNA, zaburza struktur pre-miRNA i hamuje proces wycinania miRNA z pre-miRNA. Oligomer wedBug zastrz. 1, znamienny tym, |e oligomer RNA dziaBa dwuetapowo: i) jako inhibitor kompetycyjny, na etapie wizania z rybonukleaz Dicer, ii) jako czsteczka zaburzajca struktur substratu pre-miRNA, na etapie wizania z substratem pre-miRNA. Oligomer wedBug zastrz. 1, znamienny tym, |e oligomer RNA dziaBa jednoetapowo jako czsteczka zaburzajca struktur substratu pre-miRNA, na etapie wizania z substratem pre-miRNA. Oligomer wedBug zastrz. 1, znamienny tym, |e jest wybrany spo[rd: - oligomeru RNA oddziaBujcego z rybonukleaz Dicer, - oligomeru RNA oddziaBujcego z substratem pre-miRNA, - oligomeru RNA oddziaBujcego z jednoniciowymi rejonami substratu pre-miRNA, - oligomeru RNA oddziaBujcego z dwuniciowymi rejonami substratu pre-miRNA, - oligomeru RNA oddziaBujcego z jednoniciowymi oraz dwuniciowymi rejonami substratu pre-miRNA, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA w taki sposb, i| substrat nie jest specyficznie rozpoznawany przez rybonukleaz Dicer, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA w taki sposb, i| nie dochodzi do powstawania specyficznych produktw miRNA, - oligomeru RNA zaburzajcego struktur substratu pre-miRNA w taki sposb, i| nie dochodzi do powstawania funkcjonalnych miRNA, - oligomeru RNA komplementarnego do jednoniciowych rejonw prekursora pre-miRNA, - oligomeru RNA komplementarnego do dwuniciowych rejonw prekursora pre-miRNA, - oligomeru RNA komplementarnego do jednoniciowych oraz dwuniciowych rejonw prekursora pre-miRNA, - oligomeru RNA zawierajcy modyfikacje zasad, cukru, reszt fosforanowych, przy czym oligomery s dBugo[ci do 75 nt 5. Oligomer wedBug zastrz. 1, znamienny tym, |e zawiera sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA, wybrane spo[rd sekwencji: seq. nr 2, seq. nr 3, seq. nr 4, seq. nr 8, seq. nr 9, seq.10, seq. nr 11. 5. Sposb regulowania procesu powstawania miRNA, znamienny tym, |e obejmuje etapy, w ktrych oligomer okre[lony zastrze|eniami 1 do 4 oddziaBuje z pre-miRNA, zaburza struktur pre-miRNA i hamuje proces wycinania miRNA z pre-miRNA przez ludzk rybonukleaz Dicer. 6. Sposb wedBug zastrz. 5, znamienny tym, |e stanowi wybirczy i selektywny sposb inhibicji powstawania miRNA. 7. Sposb wedBug zastrz. 5 albo 6, znamienny tym, |e oligomer dziaBa dwuetapowo: a) wi|e si z Dicer, b) produkt cicia oligomeru przez ludzk rybonukleaz Dicer wi|e si z fragmentem ptli terminalnej pre-miRNA i zmienia struktur drugo- i trzeciorzdow pre-miRNA, tak |e pre-miRNA nie jest specyficznie rozpoznawany i city przez ludzk rybonukleaz Dicer i nie dochodzi do powstawania miRNA. 8. Sposb wedBug zastrz. 5 albo 6, znamienny tym, |e oligomer dziaBa jednoetapowo: tworzy kompleks z pre-miRNA i zmienia jego struktur, tak |e pre-miRNA nie jest specyficznie rozpoznawany i city przez ludzk rybonukleaz Dicer i nie dochodzi do powstawania miRNA. 9. Sposb wedBug zastrz. 5, znamienny tym, |e sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA, wybiera si spo[rd sekwencji: seq. nr 2, seq. nr 3, seq. nr 4, seq. nr 8, seq. nr 9, seq.10, seq. nr 11. 10. Zastosowanie oligomerw RNA okre[lonych zastrze|eniami 1 do 4 zaburzajcych struktur pre-miRNA jako regulatorw procesu powstawania miRNA. 11. Zastosowanie wedBug zastrz. 10, w ktrym sekwencje komplementarne do rejonw jednoniciowych oraz dwuniciowych czsteczki pre-miRNA, wybrane s spo[rd sekwencji: seq. nr 2, seq. nr 3, seq. nr 4, seq. nr 8, seq. nr 9, seq.10, seq. nr 11. ZgBaszajcy: Instytut Chemii Bioorganicznej PAN PeBnomocnik: dr Aleksandra Twardowska rzecznik patentowy Skrt opisu Przedmiotem rozwizania s oligomer RNA, sposb regulowania procesu powstawania miRNA oraz zastosowanie oligomerw RNA jako regulatorw procesu powstawania miRNA. Bardziej szczegBowo, rozwizanie dotyczy zastosowania oligomerw RNA zaburzajcych struktur pre-miRNA jako regulatorw procesu powstawania miRNA. OddziaBywania oligomeru z prekursorem miRNA (pre-miRNA) zmieniaj struktur drugo- i trzeciorzdow prekursora. 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