{"PubmedArticle":{"MedlineCitation":{"@attributes":{"Status":"MEDLINE","Owner":"NLM","IndexingMethod":"Manual"},"PMID":{"@attributes":{"Version":"1"},"@text":"10485881"},"DateCompleted":{"Year":"1999","Month":"10","Day":"14"},"DateRevised":{"Year":"2019","Month":"05","Day":"03"},"Article":{"@attributes":{"PubModel":"Print"},"Journal":{"ISSN":{"@attributes":{"IssnType":"Print"},"@text":"0027-8424"},"JournalIssue":{"@attributes":{"CitedMedium":"Print"},"Volume":"96","Issue":"19","PubDate":{"Year":"1999","Month":"Sep","Day":"14"}},"Title":"Proceedings of the National Academy of Sciences of the United States of America","ISOAbbreviation":"Proc Natl Acad Sci U S A"},"ArticleTitle":"Evidence for the function of an exonic splicing enhancer after the first catalytic step of pre-mRNA splicing.","Pagination":{"StartPage":"10655","EndPage":"10660","MedlinePgn":"10655-60"},"Abstract":{"AbstractText":["Exonic splicing enhancers (ESEs) activate pre-mRNA splicing by promoting the use of the flanking splice sites. They are recognized by members of the serine\/arginine-rich (SR) family of proteins, such as splicing factor 2\/alternative splicing factor (SF2\/ASF), which recruit basal splicing factors to form the initial complexes during spliceosome assembly. The in vitro splicing kinetics of an ESE-dependent IgM pre-mRNA suggested that an SF2\/ASF-specific ESE has additional functions later in the splicing reaction, after the completion of the first catalytic step. A bimolecular exon ligation assay, which physically uncouples the first and second catalytic steps of splicing in a trans-splicing reaction, was adapted to test the function of the ESE after the first step. A 3' exon containing the SF2\/ASF-specific ESE underwent bimolecular exon ligation, whereas 3' exons without the ESE or with control sequences did not. The ESE-dependent trans-splicing reaction occurred after inactivation of U1 or U2 small nuclear ribonucleoprotein particles, compatible with a functional assay for events after the first step of splicing. The ESE-dependent step appears to take place before the ATP-independent part of the second catalytic step. Bimolecular exon ligation also occurred in an S100 cytosolic extract, requiring both the SF2\/ASF-dependent ESE and complementation with SF2\/ASF. These data suggest that some ESEs can act late in the splicing reaction, together with appropriate SR proteins, to enhance the second catalytic step of splicing."]},"AuthorList":{"@attributes":{"CompleteYN":"Y"},"Author":[{"@attributes":{"ValidYN":"Y"},"LastName":"Chew","ForeName":"S L","Initials":"SL","AffiliationInfo":[{"Affiliation":"Cold Spring Harbor Laboratory, Cold Spring Harbor, NY 11724, USA."}]},{"@attributes":{"ValidYN":"Y"},"LastName":"Liu","ForeName":"H X","Initials":"HX"},{"@attributes":{"ValidYN":"Y"},"LastName":"Mayeda","ForeName":"A","Initials":"A"},{"@attributes":{"ValidYN":"Y"},"LastName":"Krainer","ForeName":"A R","Initials":"AR"}]},"Language":["eng"],"GrantList":{"@attributes":{"CompleteYN":"Y"},"Grant":[{"Agency":"Wellcome Trust","Country":"United Kingdom"},{"GrantID":"R01 GM042699","Acronym":"GM","Agency":"NIGMS NIH HHS","Country":"United States"},{"GrantID":"R37 GM042699","Acronym":"GM","Agency":"NIGMS NIH HHS","Country":"United States"},{"GrantID":"GM42699","Acronym":"GM","Agency":"NIGMS NIH HHS","Country":"United States"}]},"PublicationTypeList":{"PublicationType":[{"@attributes":{"UI":"D016428"},"@text":"Journal Article"},{"@attributes":{"UI":"D013485"},"@text":"Research Support, Non-U.S. Gov't"},{"@attributes":{"UI":"D013486"},"@text":"Research Support, U.S. Gov't, Non-P.H.S."},{"@attributes":{"UI":"D013487"},"@text":"Research Support, U.S. Gov't, P.H.S."}]}},"MedlineJournalInfo":{"Country":"United States","MedlineTA":"Proc Natl Acad Sci U S A","NlmUniqueID":"7505876","ISSNLinking":"0027-8424"},"ChemicalList":{"Chemical":[{"RegistryNumber":"0","NameOfSubstance":{"@attributes":{"UI":"D007075"},"@text":"Immunoglobulin M"}},{"RegistryNumber":"0","NameOfSubstance":{"@attributes":{"UI":"D009687"},"@text":"Nuclear Proteins"}},{"RegistryNumber":"0","NameOfSubstance":{"@attributes":{"UI":"D009841"},"@text":"Oligonucleotides"}},{"RegistryNumber":"0","NameOfSubstance":{"@attributes":{"UI":"D012322"},"@text":"RNA Precursors"}},{"RegistryNumber":"0","NameOfSubstance":{"@attributes":{"UI":"D016601"},"@text":"RNA-Binding Proteins"}},{"RegistryNumber":"170974-22-8","NameOfSubstance":{"@attributes":{"UI":"D000068103"},"@text":"Serine-Arginine Splicing Factors"}},{"RegistryNumber":"8L70Q75FXE","NameOfSubstance":{"@attributes":{"UI":"D000255"},"@text":"Adenosine Triphosphate"}}]},"CitationSubset":["IM"],"MeshHeadingList":{"MeshHeading":[{"DescriptorName":{"@attributes":{"UI":"D000255","MajorTopicYN":"N"},"@text":"Adenosine Triphosphate"},"QualifierName":[{"@attributes":{"UI":"Q000494","MajorTopicYN":"N"},"@text":"pharmacology"}]},{"DescriptorName":{"@attributes":{"UI":"D000256","MajorTopicYN":"N"},"@text":"Adenoviridae"},"QualifierName":[{"@attributes":{"UI":"Q000235","MajorTopicYN":"N"},"@text":"genetics"}]},{"DescriptorName":{"@attributes":{"UI":"D002384","MajorTopicYN":"Y"},"@text":"Catalysis"}},{"DescriptorName":{"@attributes":{"UI":"D004742","MajorTopicYN":"N"},"@text":"Enhancer Elements, Genetic"},"QualifierName":[{"@attributes":{"UI":"Q000502","MajorTopicYN":"Y"},"@text":"physiology"}]},{"DescriptorName":{"@attributes":{"UI":"D004926","MajorTopicYN":"N"},"@text":"Escherichia coli"},"QualifierName":[{"@attributes":{"UI":"Q000235","MajorTopicYN":"N"},"@text":"genetics"}]},{"DescriptorName":{"@attributes":{"UI":"D005091","MajorTopicYN":"N"},"@text":"Exons"}},{"DescriptorName":{"@attributes":{"UI":"D007075","MajorTopicYN":"N"},"@text":"Immunoglobulin M"},"QualifierName":[{"@attributes":{"UI":"Q000378","MajorTopicYN":"N"},"@text":"metabolism"}]},{"DescriptorName":{"@attributes":{"UI":"D007700","MajorTopicYN":"N"},"@text":"Kinetics"}},{"DescriptorName":{"@attributes":{"UI":"D008957","MajorTopicYN":"N"},"@text":"Models, Genetic"}},{"DescriptorName":{"@attributes":{"UI":"D009687","MajorTopicYN":"N"},"@text":"Nuclear Proteins"},"QualifierName":[{"@attributes":{"UI":"Q000235","MajorTopicYN":"N"},"@text":"genetics"},{"@attributes":{"UI":"Q000378","MajorTopicYN":"N"},"@text":"metabolism"}]},{"DescriptorName":{"@attributes":{"UI":"D009841","MajorTopicYN":"N"},"@text":"Oligonucleotides"},"QualifierName":[{"@attributes":{"UI":"Q000378","MajorTopicYN":"N"},"@text":"metabolism"}]},{"DescriptorName":{"@attributes":{"UI":"D012322","MajorTopicYN":"N"},"@text":"RNA Precursors"},"QualifierName":[{"@attributes":{"UI":"Q000378","MajorTopicYN":"Y"},"@text":"metabolism"}]},{"DescriptorName":{"@attributes":{"UI":"D012326","MajorTopicYN":"N"},"@text":"RNA Splicing"},"QualifierName":[{"@attributes":{"UI":"Q000502","MajorTopicYN":"Y"},"@text":"physiology"}]},{"DescriptorName":{"@attributes":{"UI":"D016601","MajorTopicYN":"N"},"@text":"RNA-Binding Proteins"}},{"DescriptorName":{"@attributes":{"UI":"D000068103","MajorTopicYN":"N"},"@text":"Serine-Arginine Splicing 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