text:
RNA splicing
From Wikipedia, the free encyclopedia
Jump to: navigation, search
This article may be too technical for most readers to understand. Please help improve it to make it understandable to non-experts, without removing the technical details. (December 2016) (Learn how and when to remove this template message)
In molecular biology, splicing is the editing of the nascent precursor messenger RNA (pre-mRNA) transcript into a mature messenger RNA (mRNA). After splicing, introns are removed and exons are joined together (ligated). For nuclear-encoded genes, splicing takes place within the nucleus either during or immediately after transcription. For those eukaryotic genes that contain introns, splicing is usually required in order to create an mRNA molecule that can be translated into protein. For many eukaryotic introns, splicing is carried out in a series of reactions which are catalyzed by the spliceosome, a complex of snRNPs. Self-splicing introns, or ribozymes capable of catalyzing their own excision from their parent RNA molecule, also exist.
Contents
[hide]
1 Splicing pathways
1.1 Spliceosomal
1.1.1 Introns
1.1.2 Formation and activity
1.2 Self-splicing
1.3 tRNA splicing
2 Evolution
3 Biochemical mechanism
4 Alternative splicing
5 Splicing response to DNA damage
6 Experimental manipulation of splicing
7 Splicing errors and variation
8 Protein splicing
9 See also
10 References
11 External links
Splicing pathways[edit]
Several methods of RNA splicing occur in nature; the type of splicing depends on the structure of the spliced intron and the catalysts required for splicing to occur.
Spliceosomal[edit]
Introns[edit]
The word intron is derived from the term intervening sequence, that is, a segment of DNA that "intervenes" between two exons of a gene. The term intron refers to both the DNA sequence within a gene and the corresponding sequence in the unprocessed RNA transcript. As part of the RNA processing pathway, introns are removed by RNA splicing either shortly after or concurrent with transcription.[1] Introns are found in the genes of most organisms and many viruses. They can be located in a wide range of genes, including those that generate proteins, ribosomal RNA (rRNA), and transfer RNA (tRNA).[2]
Spliceosomal introns often reside within the sequence of eukaryotic protein-coding genes. Within the intron, a donor site (5' end of the intron), a branch site (near the 3' end of the intron) and an acceptor site (3' end of the intron) are required for splicing. The splice donor site includes an almost invariant sequence GU at the 5' end of the intron, within a larger, less highly conserved region. The splice acceptor site at the 3' end of the intron terminates the intron with an almost invariant AG sequence. Upstream (5'-ward) from the AG there is a region high in pyrimidines (C and U), or polypyrimidine tract. Further upstream from the polypyrimidine tract is the branchpoint, which includes an adenine nucleotide involved in lariat formation.[3][4] The consensus sequence for an intron (in IUPAC nucleic acid notation) is: G-G-[cut]-G-U-R-A-G-U (donor site) ... intron sequence ... Y-U-R-A-C (branch sequence 20-50 nucleotides upstream of acceptor site) ... Y-rich-N-C-A-G-[cut]-G (acceptor site).[5] However, it is noted that the specific sequence of intronic splicing elements and the number of nucleotides between the branchpoint and the nearest 3’ acceptor site affect splice site selection.[6][7] Also, point mutations in the underlying DNA or errors during transcription can activate a cryptic splice site in part of the transcript that usually is not spliced. This results in a mature messenger RNA with a missing section of an exon. In this way, a point mutation, which might otherwise affect only a single amino acid, can manifest as a deletion or truncation in the final protein.
Simple illustration of exons and introns in pre-mRNA
Formation and activity[edit]
Splicing is catalyzed by the spliceosome, a large RNA-protein complex composed of five small nuclear ribonucleoproteins (snRNPs, pronounced 'snurps'). Assembly and activity of the spliceosome occurs during transcription of the pre-mRNA. The RNA components of snRNPs interact with the intron and are involved in catalysis. Two types of spliceosomes have been identified (major and minor) which contain different snRNPs.
The major spliceosome splices introns containing GU at the 5' splice site and AG at the 3' splice site. It is composed of the U1, U2, U4, U5, and U6 snRNPs and is active in the nucleus. In addition, a number of proteins including U2 small nuclear RNA auxiliary factor 1 (U2AF35), U2AF2 (U2AF65)[8] and SF1 are required for the assembly of the spliceosome.[4][9] The spliceosome forms different complexes during the splicing process:[10]
Complex E
The U1 snRNP binds to the GU sequence at the 5' splice site of an intron;
Splicing factor 1 binds to the intron branch point sequence;
U2AF1 binds at the 3' splice site of the intron;
U2AF2 binds to the polypyrimidine tract;[11]
Complex A (pre-spliceosome)
The U2 snRNP displaces SF1 and binds to the branch point sequence and ATP is hydrolyzed;
Complex B (pre-catalytic spliceosome)
The U5/U4/U6 snRNP trimer binds, and the U5 snRNP binds exons at the 5' site, with U6 binding to U2;
Complex B*
The U1 snRNP is released, U5 shifts from exon to intron, and the U6 binds at the 5' splice site;
Complex C (catalytic spliceosome)
U4 is released, U6/U2 catalyzes transesterification, making the 5'-end of the intron ligate to the A on intron and form a lariat, U5 binds exon at 3' splice site, and the 5' site is cleaved, resulting in the formation of the lariat;
Complex C* (post-spliceosomal complex)
U2/U5/U6 remain bound to the lariat, and the 3' site is cleaved and exons are ligated using ATP hydrolysis. The spliced RNA is released, the lariat is released and degraded,[12] and the snRNPs are recycled.
This type of splicing is termed canonical splicing or termed the lariat pathway, which accounts for more than 99% of splicing. By contrast, when the intronic flanking sequences do not follow the GU-AG rule, noncanonical splicing is said to occur (see "minor spliceosome" below).[13]
The minor spliceosome is very similar to the major spliceosome, but instead it splices out rare introns with different splice site sequences. While the minor and major spliceosomes contain the same U5 snRNP, the minor spliceosome has different but functionally analogous snRNPs for U1, U2, U4, and U6, which are respectively called U11, U12, U4atac, and U6atac.[14] Unlike the major spliceosome, it is found outside the nucleus, but very close to the nuclear membrane.[15]
Trans-splicing is a form of splicing that joins two exons that are not within the same RNA transcript.[16]
Self-splicing[edit]
Self-splicing occurs for rare introns that form a ribozyme, performing the functions of the spliceosome by RNA alone. There are three kinds of self-splicing introns, Group I, Group II and Group III. Group I and II introns perform splicing similar to the spliceosome without requiring any protein. This similarity suggests that Group I and II introns may be evolutionarily related to the spliceosome. Self-splicing may also be very ancient, and may have existed in an RNA world present before protein.
Two transesterifications characterize the mechanism in which group I introns are spliced:
3'OH of a free guanine nucleoside (or one located in the intron) or a nucleotide cofactor (GMP, GDP, GTP) attacks phosphate at the 5' splice site.
3'OH of the 5' exon becomes a nucleophile and the second transesterification results in the joining of the two exons.
The mechanism in which group II introns are spliced (two transesterification reaction like group I introns) is as follows:
The 2'OH of a specific adenosine in the intron attacks the 5' splice site, thereby forming the lariat
The 3'OH of the 5' exon triggers the second transesterification at the 3' splice site, thereby joining the exons together.
tRNA splicing[edit]
tRNA (also tRNA-like) splicing is another rare form of splicing that usually occurs in tRNA. The splicing reaction involves a different biochemistry than the spliceosomal and self-splicing pathways.
In the yeast Saccharomyces cerevisiae, a yeast tRNA splicing endonuclease heterotetramer, composed of TSEN54, TSEN2, TSEN34, and TSEN15, cleaves pre-tRNA at two sites in the acceptor loop to form a 5'-half tRNA, terminating at a 2',3'-cyclic phosphodiester group, and a 3'-half tRNA, terminating at a 5'-hydroxyl group, along with a discarded intron.[17] Yeast tRNA kinase then phosphorylates the 5'-hydroxyl group using adenosine triphosphate. Yeast tRNA cyclic phosphodiesterase cleaves the cyclic phosphodiester group to form a 2'-phosphorylated 3' end. Yeast tRNA ligase adds an adenosine monophosphate group to the 5' end of the 3'-half and joins the two halves together.[18] NAD-dependent 2'-phosphotransferase then removes the 2'-phosphate group.[19][20]
Evolution[edit]
Splicing occurs in all the kingdoms or domains of life, however, the extent and types of splicing can be very different between the major divisions. Eukaryotes splice many protein-coding messenger RNAs and some non-coding RNAs. Prokaryotes, on the other hand, splice rarely and mostly non-coding RNAs. Another important difference between these two groups of organisms is that prokaryotes completely lack the spliceosomal pathway.
Because spliceosomal introns are not conserved in all species, there is debate concerning when spliceosomal splicing evolved. Two models have been proposed: the intron late and intron early models (see intron evolution).
Splicing diversity
Eukaryotes
Prokaryotes
Spliceosomal + −
Self-splicing + +
tRNA + +
Biochemical mechanism[edit]
Diagram illustrating the two-step biochemistry of splicing
Spliceosomal splicing and self-splicing involve a two-step biochemical process. Both steps involve transesterification reactions that occur between RNA nucleotides. tRNA splicing, however, is an exception and does not occur by transesterification.[21]
Spliceosomal and self-splicing transesterification reactions occur via two sequential transesterification reactions. First, the 2'OH of a specific branchpoint nucleotide within the intron, defined during spliceosome assembly, performs a nucleophilic attack on the first nucleotide of the intron at the 5' splice site, forming the lariat intermediate. Second, the 3'OH of the released 5' exon then performs a nucleophilic attack at the first nucleotide following the last nucleotide of the intron at the 3' splice site, thus joining the exons and releasing the intron lariat.[22]
Alternative splicing[edit]
Main article: Alternative splicing
In many cases, the splicing process can create a range of unique proteins by varying the exon composition of the same mRNA. This phenomenon is then called alternative splicing. Alternative splicing can occur in many ways. Exons can be extended or skipped, or introns can be retained. It is estimated that 95% of transcripts from multiexon genes undergo alternative splicing, some instances of which occur in a tissue-specific manner and/or under specific cellular conditions.[23] Development of high throughput mRNA sequencing technology can help quantify the expression levels of alternatively spliced isoforms. Differential expression levels across tissues and cell lineages allowed computational approaches to be developed to predict the functions of these isoforms.[24][25] Given this complexity, alternative splicing of pre-mRNA transcripts is regulated by a system of trans-acting proteins (activators and repressors) that bind to cis-acting sites or "elements" (enhancers and silencers) on the pre-mRNA transcript itself. These proteins and their respective binding elements promote or reduce the usage of a particular splice site. The binding specificity comes from the sequence and structure of the cis-elements, e.g. in HIV-1 there are many donor and acceptor splice sites. Among the various splice sites, ssA7, which is 3' acceptor site, folds into three stem loop structures, i.e. Intronic splicing silencer (ISS), Exonic splicing enhancer (ESE), and Exonic splicing silencer (ESSE3). Solution structure of Intronic splicing silencer and its interaction to host protein hnRNPA1 give insight into specific recognition.[26] However, adding to the complexity of alternative splicing, it is noted that the effects of regulatory factors are many times position-dependent. For example, a splicing factor that serves as a splicing activator when bound to an intronic enhancer element may serve as a repressor when bound to its splicing element in the context of an exon, and vice versa.[27] In addition to the position-dependent effects of enhancer and silencer elements, the location of the branchpoint (i.e., distance upstream of the nearest 3’ acceptor site) also affects splicing.[6] The secondary structure of the pre-mRNA transcript also plays a role in regulating splicing, such as by bringing together splicing elements or by masking a sequence that would otherwise serve as a binding element for a splicing factor.[28][29]
Splicing response to DNA damage[edit]
DNA damage affects splicing factors by altering their post-translational modification, localization, expression and activity.[30] Furthermore, DNA damage often disrupts splicing by interfering with its coupling to transcription. DNA damage also has an impact on the splicing and alternative splicing of genes intimately associated with DNA repair.[30] For instance, DNA damages modulate the alternative splicing of the DNA repair genes Brca1 and Ercc1.
Experimental manipulation of splicing[edit]
Splicing events can be experimentally altered[31][32] by binding steric-blocking antisense oligos such as Morpholinos or Peptide nucleic acids to snRNP binding sites, to the branchpoint nucleotide that closes the lariat,[33] or to splice-regulatory element binding sites.[34]
Splicing errors and variation[edit]
It has been suggested that one third of all disease-causing mutations impact on splicing.[27] Common errors include:
Mutation of a splice site resulting in loss of function of that site. Results in exposure of a premature stop codon, loss of an exon, or inclusion of an intron.
Mutation of a splice site reducing specificity. May result in variation in the splice location, causing insertion or deletion of amino acids, or most likely, a disruption of the reading frame.
Displacement of a splice site, leading to inclusion or exclusion of more RNA than expected, resulting in longer or shorter exons.
Although many splicing errors are safeguarded by a cellular quality control mechanism termed nonsense-mediated mRNA decay (NMD),[35] a number of splicing-related diseases also exist, as suggested above.[36]
Allelic differences in mRNA splicing are likely to be a common and important source of phenotypic diversity at the molecular level, in addition to their contribution to genetic disease susceptibility. Indeed, genome-wide studies in humans have identified a range of genes that are subject to allele-specific splicing.
Protein splicing[edit]
Main article: Protein splicing
In addition to RNA, proteins can undergo splicing. Although the biomolecular mechanisms are different, the principle is the same: parts of the protein, called inteins instead of introns, are removed. The remaining parts, called exteins instead of exons, are fused together. Protein splicing has been observed in a wide range of organisms, including bacteria, archaea, plants, yeast and humans.[37]
See also[edit]
Wikimedia Commons has media related to Splicing.
cDNA
post-transcriptional modification
mRNA capping
polyadenylation
Exon Junction Complex
SWAP protein domain, a splicing regulator
References[edit]
Jump up ^ Tilgner, Hagen; Knowles, David G.; Johnson, Rory; Davis, Carrie A.; Chakrabortty, udipto; Djebali, Sarah; Curado, João; Snyder, Michael; Gingeras, Thomas R.; Guigó, Roderic (2012). "Deep sequencing of subcellular RNA fractions shows splicing to be predominantly co-transcriptional in the human genome but inefficient for lncRNAs". Genome Research. 22 (9): 1616–25. doi:10.1101/gr.134445.111. PMC 3431479  . PMID 22955974.
Jump up ^ William Roy, Scott; Gilbert, Walter (2006). "The evolution of spliceosomal introns: Patterns, puzzles and progress". Nature Reviews Genetics. 7 (3): 211–21. doi:10.1038/nrg1807. PMID 16485020.
Jump up ^ Clancy, Suzanne (2008). "RNA Splicing: Introns, Exons and Spliceosome". Nature Education. 1 (1): 31. Retrieved 31 March 2011.
^ Jump up to: a b Black, Douglas L. (June 2003). "Mechanisms of Alternative Pre-Messenger RNA Splicing". Annual Review of Biochemistry. 72 (1): 291–336. doi:10.1146/annurev.biochem.72.121801.161720. PMID 12626338.
Jump up ^ "Molecular Biology of the Cell". 2012 Journal Citation Reports. Web of Science (Science ed.). Thomson Reuters. 2013.
^ Jump up to: a b Taggart, Allison J; Desimone, Alec M; Shih, Janice S; Filloux, Madeleine E; Fairbrother, William G (2012). "Large-scale mapping of branchpoints in human pre-mRNA transcripts in vivo". Nature Structural & Molecular Biology. 19 (7): 719–21. doi:10.1038/nsmb.2327. PMC 3465671  . PMID 22705790.
Jump up ^ Corvelo, André; Hallegger, Martina; Smith, Christopher W. J.; Eyras, Eduardo (2010). Meyer, Irmtraud M, ed. "Genome-Wide Association between Branch Point Properties and Alternative Splicing". PLoS Computational Biology. 6 (11): e1001016. doi:10.1371/journal.pcbi.1001016. PMC 2991248  . PMID 21124863.
Jump up ^ Graveley, Brenton R; Hertel, Klemens J; Maniatis, Tom (June 2001). "The role of U2AF35 and U2AF65 in enhancer-dependent splicing". RNA. 7 (6): 806–818. doi:10.1017/s1355838201010317. PMC 1370132  . PMID 11421359. Retrieved 17 December 2014.
Jump up ^ Matlin, Arianne J.; Clark, Francis; Smith, Christopher W. J. (2005). "Understanding alternative splicing: Towards a cellular code". Nature Reviews Molecular Cell Biology. 6 (5): 386–98. doi:10.1038/nrm1645. PMID 15956978.
Jump up ^ Matera, A. Gregory; Wang, Zefeng (23 January 2014). "A day in the life of the spliceosome". Nature Reviews Molecular Cell Biology. 15 (2): 108–121. doi:10.1038/nrm3742. PMC 4060434  . PMID 24452469. Retrieved 12 December 2014.
Jump up ^ Guth, Sabine; Valcárcel, Juan (1 December 2000). "Kinetic role for mammalian SF1/BBP in spliceosome assembly and function after polypyrimidine tract recognition by U2AF". Journal of Biological Chemistry. 275 (48): 38059–66. doi:10.1074/jbc.M001483200. PMID 10954700. Retrieved 12 December 2014.
Jump up ^ Cheng, Zhi; Menees, Thomas M. (8 November 2011). "RNA splicing and debranching viewed through analysis of RNA lariats". Molecular Genetics and Genomics. 286 (5–6): 395–410. doi:10.1007/s00438-011-0635-y. Retrieved 12 December 2014.
Jump up ^ Ng, Bernard; Yang, Fan; Huston, David P.; Yan, Yan; Yang, Yu; Xiong, Zeyu; Peterson, Leif E.; Wang, Hong; Yang, Xiao-Feng (2004). "Increased noncanonical splicing of autoantigen transcripts provides the structural basis for expression of untolerized epitopes". Journal of Allergy and Clinical Immunology. 114 (6): 1463–70. doi:10.1016/j.jaci.2004.09.006. PMC 3902068  . PMID 15577853.
Jump up ^ Patel, Abhijit A.; Steitz, Joan A. (2003). "Splicing double: Insights from the second spliceosome". Nature Reviews Molecular Cell Biology. 4 (12): 960–70. doi:10.1038/nrm1259. PMID 14685174.
Jump up ^ König, Harald; Matter, Nathalie; Bader, Rüdiger; Thiele, Wilko; Müller, Ferenc (2007). "Splicing Segregation: The Minor Spliceosome Acts outside the Nucleus and Controls Cell Proliferation". Cell. 131 (4): 718–29. doi:10.1016/j.cell.2007.09.043. PMID 18022366.
Jump up ^ Di Segni, Gianfranco; Gastaldi, Serena; Tocchini-Valentini, Glauco P. (2008). "Cis- and trans-splicing of mRNAs mediated by tRNA sequences in eukaryotic cells". Proceedings of the National Academy of Sciences. 105 (19): 6864–9. Bibcode:2008PNAS..105.6864D. doi:10.1073/pnas.0800420105. JSTOR 25461891. PMC 2383978  . PMID 18458335.
Jump up ^ Trotta, Christopher R; Miao, Feng; Arn, Eric A; Stevens, Scott W; Ho, Calvin K; Rauhut, Reinhard; Abelson, John N (June 1997). "The Yeast tRNA Splicing Endonuclease: A Tetrameric Enzyme with Two Active Site Subunits Homologous to the Archaeal tRNA Endonucleases". Cell. 89 (6): 849–858. doi:10.1016/S0092-8674(00)80270-6. Retrieved 17 December 2014.
Jump up ^ Westaway, Shawn K; Phizicky, Eric M; Abelson, John (5 March 1988). "Structure and function of the yeast tRNA ligase gene". The Journal of Biological Chemistry. 263 (7): 3171–6. PMID 3277966. Retrieved 17 December 2014.
Jump up ^ Paushkin, Sergey V; Patel, Meenal; Furia, Bansri S; Peltz, Stuart W; Trotta, Christopher R (April 2004). "Identification of a Human Endonuclease Complex Reveals a Link between tRNA Splicing and Pre-mRNA 3′ End Formation". Cell. 117 (3): 311–321. doi:10.1016/S0092-8674(04)00342-3. PMID 15109492. Retrieved 17 December 2014.
Jump up ^ Soma, Akiko (1 April 2014). "Circularly permuted tRNA genes: their expression and implications for their physiological relevance and development". Frontiers in Genetics. 5. doi:10.3389/fgene.2014.00063. Retrieved 17 December 2014.
Jump up ^ Abelson, John; Trotta, Christopher R; Li, Hong (22 May 1998). "tRNA Splicing". Journal of Biological Chemistry. 273 (21): 12685–12688. doi:10.1074/jbc.273.21.12685. PMID 9582290. Retrieved 17 December 2014.
Jump up ^ Fica, Sebastian M.; Tuttle, Nicole; Novak, Thaddeus; Li, Nan-Sheng; Lu, Jun; Koodathingal, Prakash; Dai, Qing; Staley, Jonathan P.; Piccirilli, Joseph A. (6 November 2013). "RNA catalyses nuclear pre-mRNA splicing". Nature. 503 (7475): 229–234. doi:10.1038/nature12734. PMC 4666680  . PMID 24196718. Retrieved 17 December 2014.
Jump up ^ Pan, Qun; Shai, Ofer; Lee, Leo J; Frey, Brendan J; Blencowe, Benjamin J (2008). "Deep surveying of alternative splicing complexity in the human transcriptome by high-throughput sequencing". Nature Genetics. 40 (12): 1413–5. doi:10.1038/ng.259. PMID 18978789.
Jump up ^ Eksi, R; Li, HD; Menon, R; Wen, Y; Omenn, GS; Kretzler, M; Guan, Y (Nov 2013). "Systematically differentiating functions for alternatively spliced isoforms through integrating RNA-seq data". PLOS Computational Biology. 9 (11): e1003314. doi:10.1371/journal.pcbi.1003314. PMC 3820534  . PMID 24244129.
Jump up ^ Li, HD; Menon, R; Omenn, GS; Guan, Y (Jun 17, 2014). "The emerging era of genomic data integration for analyzing splice isoform function". Trends in genetics : TIG. 30 (8): 340–347. doi:10.1016/j.tig.2014.05.005. PMC 4112133  . PMID 24951248.
Jump up ^ Jain, Niyati; Morgan, Christopher; Rife, Brittney; Salemi, Marco; Tolbert, Blanton (24 November 2015). "Solution Structure of the HIV-1 Intron Splicing Silencer and its Interactions with the UP1 Domain of hnRNP A1". Journal of Biological Sciences.
^ Jump up to: a b Lim, Kian Huat; Ferraris, Luciana; Filloux, Madeleine E.; Raphael, Benjamin J.; Fairbrother, William G. (2011). "Using positional distribution to identify splicing elements and predict pre-mRNA processing defects in human genes". Proceedings of the National Academy of Sciences. 108 (27): 11093–8. doi:10.1073/pnas.1101135108. PMC 3131313  . PMID 21685335.
Jump up ^ Warf, M. Bryan; Berglund, J. Andrew (2010). "Role of RNA structure in regulating pre-mRNA splicing". Trends in Biochemical Sciences. 35 (3): 169–78. doi:10.1016/j.tibs.2009.10.004. PMC 2834840  . PMID 19959365.
Jump up ^ Reid, D. C.; Chang, B. L.; Gunderson, S. I.; Alpert, L.; Thompson, W. A.; Fairbrother, W. G. (2009). "Next-generation SELEX identifies sequence and structural determinants of splicing factor binding in human pre-mRNA sequence". RNA. 15 (12): 2385–97. doi:10.1261/rna.1821809. PMC 2779669  . PMID 19861426.
^ Jump up to: a b Shkreta L, Chabot B (2015). "The RNA Splicing Response to DNA Damage". Biomolecules. 5 (4): 2935–77. doi:10.3390/biom5042935. PMC 4693264  . PMID 26529031.
Jump up ^ Draper, Bruce W.; Morcos, Paul A.; Kimmel, Charles B. (2001). "Inhibition of zebrafishfgf8 pre-mRNA splicing with morpholino oligos: A quantifiable method for gene knockdown". Genesis. 30 (3): 154–6. doi:10.1002/gene.1053. PMID 11477696.
Jump up ^ Sazani, Peter; Kang, Shin-Hong; Maier, Martin A.; Wei, Changfu; Dillman, Jennifer; Summerton, James; Manoharan, Muthiah; Kole, Ryszard (2001). "Nuclear antisense effects of neutral, anionic and cationic oligonucleotide analogs". Nucleic Acids Research. 29 (19): 3965–74. doi:10.1093/nar/29.19.3965. PMC 60237  . PMID 11574678.
Jump up ^ Morcos, Paul A. (2007). "Achieving targeted and quantifiable alteration of mRNA splicing with Morpholino oligos". Biochemical and Biophysical Research Communications. 358 (2): 521–7. doi:10.1016/j.bbrc.2007.04.172. PMID 17493584.
Jump up ^ Bruno, I. G.; Jin, W; Cote, GJ (2004). "Correction of aberrant FGFR1 alternative RNA splicing through targeting of intronic regulatory elements". Human Molecular Genetics. 13 (20): 2409–20. doi:10.1093/hmg/ddh272. PMID 15333583.
Jump up ^ Danckwardt, S.; Neu-Yilik, G; Thermann, R; Frede, U; Hentze, MW; Kulozik, AE (2002). "Abnormally spliced beta -globin mRNAs: A single point mutation generates transcripts sensitive and insensitive to nonsense-mediated mRNA decay". Blood. 99 (5): 1811–6. doi:10.1182/blood.V99.5.1811. PMID 11861299.
Jump up ^ Ward, Amanda J; Cooper, Thomas A (2009). "The pathobiology of splicing". The Journal of Pathology. 220 (2): 152–63. doi:10.1002/path.2649. PMC 2855871  . PMID 19918805.
Jump up ^ Hanada, Ken-Ichi; Yang, James C. (2005). "Novel biochemistry: Post-translational protein splicing and other lessons from the school of antigen processing". Journal of Molecular Medicine. 83 (6): 420–8. doi:10.1007/s00109-005-0652-6. PMID 15759099.
External links[edit]
Virtual Cell Animation Collection: mRNA Splicing
RNA Splicing at the US National Library of Medicine Medical Subject Headings (MeSH)
[show]
v
t
e
Gene expression
Introduction
to genetics
Genetic code
Central dogma
DNA → RNA → Protein
Special transfers
RNA→RNA
RNA→DNA
Protein→Protein
Transcription
Types
Bacterial
Eukaryotic
Key elements
Transcription factor
RNA polymerase
Promoter
Post-transcription
Precursor mRNA (pre-mRNA / hnRNA)
5' capping
Splicing
Polyadenylation
Histone acetylation and deacetylation
Translation
Types
Prokaryotic
Eukaryotic
Key elements
Ribosome
Transfer RNA (tRNA)
Ribosome-nascent chain complex (RNC)
Post-translational modification (functional groups · peptides · structural changes)
Regulation
Epigenetic
imprinting
Transcriptional
Gene regulatory network
cis-regulatory element
lac operon
Post-transcriptional
sequestration (P-bodies)
alternative splicing
microRNA
Translational
Post-translational
reversible
irreversible
Influential people
François Jacob
Jacques Monod
[show]
v
t
e
Post-transcriptional modification
Nuclear
Precursor mRNA
5′ cap formation
Polyadenylation
CPSF
CstF
PAP
PAB2
CFI
CFII
Poly(A)-binding protein
RNA editing
Polyuridylation
RNA splicing
Intron / Exon
snRNP
Spliceosome
minor
U1
Alternative splicing
pre-mRNA factors
PLRG1
PRPF3
PRPF4
PRPF4B
PRPF6
PRPF8
PRPF18
PRPF19
PRPF31
PRPF38A
PRPF38B
PRPF39
PRPF40A
PRPF40B
Cytosolic
5′ cap methylation
mRNA decapping
DCP1A
DCP1B
DCP2
DCPS
EDC3
EDC4
Retrieved from "https://en.wikipedia.org/w/index.php?title=RNA_splicing&oldid=828611016"
Categories:
Gene expression
Spliceosome
RNA splicing
Hidden categories:
Wikipedia articles that are too technical from December 2016
All articles that are too technical
Articles needing expert attention from December 2016
All articles needing expert attention
Navigation menu
Personal tools
Not logged in
Talk
Contributions
Create account
Log in
Namespaces
Article
Talk
Variants
Views
Read
Edit
View history
More
Search
Navigation
Main page
Contents
Featured content
Current events
Random article
Donate to Wikipedia
Wikipedia store
Interaction
Help
About Wikipedia
Community portal
Recent changes
Contact page
Tools
What links here
Related changes
Upload file
Special pages
Permanent link
Page information
Wikidata item
Cite this page
Print/export
Create a book
Download as PDF
Printable version
In other projects
Wikimedia Commons
Languages
Aragonés
Bosanski
Català
Čeština
Deutsch
Eesti
Ελληνικά
Español
Esperanto
فارسی
Français
Galego
Italiano
עברית
ქართული
Қазақша
Nederlands
日本語
Norsk
Polski
Português
Română
Русский
Simple English
Српски / srpski
Srpskohrvatski / српскохрватски
Suomi
Svenska
தமிழ்
Türkçe
Українська
中文
23 more
Edit links
This page was last edited on 3 March 2018, at 17:25.
Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation, Inc., a non-profit organization.
Privacy policy
About Wikipedia
Disclaimers
Contact Wikipedia
Developers
Cookie statement
Mobile view
Enable previews

key: html

type: local

value: div

attrs: id class

values: mw-page-base noprint

values: mw-head-base noprint

value: a

attrs: id class lang

values: firstHeading firstHeading en

values: siteSub noprint

values: jump-to-nav mw-jump

value: img

attrs: alt src width height srcset data-file-width data-file-height

value: b

attrs: href title

values: Wikipedia:Make technical articles understandable

value: i

value: small

value: tbody

value: table

attrs: class role

values: plainlinks metadata ambox ambox-style ambox-technical

values:
/wiki/Precursor_messenger_RNA
mw-redirect
Precursor messenger RNA

values:
/wiki/Nuclear_genes
mw-redirect
Nuclear genes

values:
/wiki/Self-splicing_intron
mw-redirect
Self-splicing intron

values:
/wiki/Transcription_(genetics)
mw-redirect
Transcription (genetics)

attrs: alt src width height class srcset data-file-width data-file-height

values:
/wiki/Nucleophilic_attack
mw-redirect
Nucleophilic attack

values: Edit section: Splicing response to DNA damage

values: Edit section: Experimental manipulation of splicing

values: Edit section: Splicing errors and variation

values:
/wiki/Nonsense-mediated_mRNA_decay
mw-redirect
Nonsense-mediated mRNA decay

values:
/wiki/Exon_Junction_Complex
mw-redirect
Exon Junction Complex

values:
nofollow
external text
//doi.org/10.1101%2Fgr.134445.111

attrs: alt src title width height srcset data-file-width data-file-height

values:
/wiki/PubMed_Identifier
mw-redirect
PubMed Identifier

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/22955974

values:
nofollow
external text
//doi.org/10.1038%2Fnrg1807

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/16485020

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/12626338

values:
nofollow
external text
//doi.org/10.1038%2Fnsmb.2327

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/22705790

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/21124863

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11421359

values:
nofollow
external text
//doi.org/10.1038%2Fnrm1645

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15956978

values:
nofollow
external text
//doi.org/10.1038%2Fnrm3742

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24452469

values:
nofollow
external text
//doi.org/10.1074%2Fjbc.M001483200

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10954700

values:
nofollow
external text
//doi.org/10.1007%2Fs00438-011-0635-y

values:
nofollow
external text
//doi.org/10.1016%2Fj.jaci.2004.09.006

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15577853

values:
nofollow
external text
//doi.org/10.1038%2Fnrm1259

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/14685174

values:
nofollow
external text
//doi.org/10.1016%2Fj.cell.2007.09.043

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/18022366

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.0800420105

values:
nofollow
external text
//www.jstor.org/stable/25461891

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/18458335

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/3277966

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15109492

values:
nofollow
external text
//doi.org/10.3389%2Ffgene.2014.00063

values:
nofollow
external text
//doi.org/10.1074%2Fjbc.273.21.12685

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/9582290

values:
nofollow
external text
//doi.org/10.1038%2Fnature12734

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24196718

values:
nofollow
external text
//doi.org/10.1038%2Fng.259

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/18978789

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24244129

values:
nofollow
external text
//doi.org/10.1016%2Fj.tig.2014.05.005

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24951248

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.1101135108

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/21685335

values:
nofollow
external text
//doi.org/10.1016%2Fj.tibs.2009.10.004

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/19959365

values:
nofollow
external text
//doi.org/10.1261%2Frna.1821809

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/19861426

values:
nofollow
external text
//doi.org/10.3390%2Fbiom5042935

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/26529031

values:
nofollow
external text
//doi.org/10.1002%2Fgene.1053

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11477696

values:
nofollow
external text
//doi.org/10.1093%2Fnar%2F29.19.3965

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11574678

values:
nofollow
external text
//doi.org/10.1016%2Fj.bbrc.2007.04.172

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/17493584

values:
nofollow
external text
//doi.org/10.1093%2Fhmg%2Fddh272

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15333583

values:
nofollow
external text
//doi.org/10.1182%2Fblood.V99.5.1811

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11861299

values:
nofollow
external text
//doi.org/10.1002%2Fpath.2649

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/19918805

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15759099

values:
/wiki/Reverse_transcription
mw-redirect
Reverse transcription

values:
row
navbox-group
width:8.0em;font-weight: normal;

values:
/wiki/Posttranslational_modification
mw-redirect
Posttranslational modification

values: nowraplinks collapsible autocollapse navbox-inner

values: Template talk:Post transcriptional modification (page does not exist)

values: Cleavage and polyadenylation specificity factor

values: nowraplinks hlist collapsible autocollapse navbox-inner

values: Category:Wikipedia articles that are too technical from December 2016

values: Category:All articles that are too technical

values: Category:Articles needing expert attention from December 2016

values: Category:All articles needing expert attention

values: Discussion about edits from this IP address [ctrl-alt-n]

values: A list of edits made from this IP address [ctrl-alt-y]

values: You are encouraged to create an account and log in; however, it is not mandatory

values: You're encouraged to log in; however, it's not mandatory. [ctrl-alt-o]

values:
/wiki/RNA_splicing
View the content page [ctrl-alt-c]
c

values: Discussion about the content page [ctrl-alt-t]

values: p-namespaces navigation vectorTabs p-namespaces-label

values:
p-variants
navigation
vectorMenu emptyPortlet
p-variants-label

values: Past revisions of this page [ctrl-alt-h]

values: p-views navigation vectorTabs p-views-label

values:
p-cactions
navigation
vectorMenu emptyPortlet
p-cactions-label

values:
mw-wiki-logo
/wiki/Main_Page
Visit the main page

values:
/wiki/Main_Page
Visit the main page [ctrl-alt-z]
z

values: Featured content – the best of Wikipedia

values: Find background information on current events

values:
/wiki/Special:Random
Load a random article [ctrl-alt-x]
x

values:
interlanguage-link interwiki-an
display: none;

values:
interlanguage-link interwiki-bs
display: none;

values:
interlanguage-link interwiki-ca
display: none;

values:
https://cs.wikipedia.org/wiki/Splicing
Splicing – Czech
cs
cs
interlanguage-link-target

values:
interlanguage-link interwiki-cs
display: none;

values:
interlanguage-link interwiki-et
display: none;

values: Συναρμογή (βιολογία) – Greek

values:
interlanguage-link interwiki-el
display: none;

values:
https://es.wikipedia.org/wiki/Splicing
Splicing – Spanish
es
es
interlanguage-link-target

values:
https://eo.wikipedia.org/wiki/Splisado
Splisado – Esperanto
eo
eo
interlanguage-link-target

values:
interlanguage-link interwiki-eo
display: none;

values: پیرایش آران‌ای – Persian

values:
interlanguage-link interwiki-fa
display: none;

values:
https://gl.wikipedia.org/wiki/Splicing
Splicing – Galician
gl
gl
interlanguage-link-target

values:
interlanguage-link interwiki-gl
display: none;

values:
https://it.wikipedia.org/wiki/Splicing
Splicing – Italian
it
it
interlanguage-link-target

values:
interlanguage-link interwiki-it
display: none;

values: שחבור – Hebrew

values:
interlanguage-link interwiki-he
display: none;

values: სპლაისინგი – Georgian

values:
interlanguage-link interwiki-ka
display: none;

values: Сплайсинг – Kazakh

values:
interlanguage-link interwiki-kk
display: none;

values:
https://nl.wikipedia.org/wiki/Splicing
Splicing – Dutch
nl
nl
interlanguage-link-target

values:
interlanguage-link interwiki-nl
display: none;

values: スプライシング – Japanese

values:
interlanguage-link interwiki-no
display: none;

values:
https://pl.wikipedia.org/wiki/Splicing
Splicing – Polish
pl
pl
interlanguage-link-target

values:
interlanguage-link interwiki-pl
display: none;

values:
https://pt.wikipedia.org/wiki/Splicing
Splicing – Portuguese
pt
pt
interlanguage-link-target

values:
interlanguage-link interwiki-ro
display: none;

values: Сплайсинг – Russian

values:
interlanguage-link interwiki-simple
display: none;

values:
interlanguage-link interwiki-sr
display: none;

values:
interlanguage-link interwiki-sh
display: none;

values:
interlanguage-link interwiki-fi
display: none;

values:
interlanguage-link interwiki-sv
display: none;

values: ஆர்.என்.ஏ முதிர்வாக்கம் – Tamil

values: Сплайсинг – Ukrainian

values:
interlanguage-link interwiki-uk
display: none;

values: All languages (initial selection from common choices by you and others)

values: /static/images/wikimedia-button-1.5x.png 1.5x, /static/images/wikimedia-button-2x.png 2x

values: /static/images/poweredby_mediawiki_132x47.png 1.5x, /static/images/poweredby_mediawiki_176x62.png 2x

values:
suggestions
display: none; font-size: 13px;

values: mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject page-RNA_splicing rootpage-RNA_splicing skin-vector action-view


text:
1967 European Cup Final
From Wikipedia, the free encyclopedia
Jump to: navigation, search
1967 European Cup Final
Match programme cover
Event
1966–67 European Cup
Celtic
Internazionale
2
1
Date
25 May 1967
Venue
Estádio Nacional, Lisbon
Referee
Kurt Tschenscher
(West Germany)
Attendance
45,000
← 1966
1968 →
The 1967 European Cup Final was a football match between Italian team Internazionale and Scottish team Celtic. It took place at the Estádio Nacional in Lisbon, Portugal on 25 May 1967 in front of a crowd of 45,000. It was the final of the 1966–67 European Cup, the premier club competition in Europe. The match was Celtic's first European final and Internazionale's third; they had won the tournament in two of the previous three years.
Both teams had to go through four qualifying rounds to get the final. Celtic won their first two ties comfortably, with their second two rounds being tighter. Internazionale's first tie was very close but they won their next two by bigger margins. In the semi-final, Internazionale needed a replay to win the tie.
Internazionale scored after seven minutes, when Sandro Mazzola converted a penalty. Celtic equalised through Tommy Gemmell after he scored on 63 minutes. Stevie Chalmers then put Celtic in the lead after 84 minutes. The match finished 2–1 to Celtic. It was said to be a victory for football because Celtic's attacking football overcame Internazionale's "catenaccio" defensive style, which was considered to be a less attractive way to play the game. Celtic's manager Jock Stein and the team received acclaim after the match and were given the nickname the Lisbon Lions; considered to be the greatest side in the club's history.
Contents
[hide]
1 Route to the final
1.1 Celtic
1.2 Internazionale
2 Match
2.1 Background
2.2 Summary
2.3 Details
3 Post-match
4 See also
5 References
Route to the final[edit]
Main article: 1966–67 European Cup
Celtic[edit]
Round
Opponents
First leg
Second leg
Aggregate score
First round
Zürich 2–0 (h) 3–0 (a) 5–0
Second round
Nantes 3–1 (a) 3–1 (h) 6–2
Quarter-finals
Vojvodina 0–1 (a) 2–0 (h) 2–1
Semi-finals
Dukla Prague 3–1 (h) 0–0 (a) 3–1
Celtic qualified for the European Cup after winning the 1965–66 Scottish First Division, their 21st title, by two points over rivals Rangers.[1] Celtic entered at the first round where they faced Swiss side Zürich. Celtic won 2–0 at home, with goals from Tommy Gemmell and Joe McBride. They then won the away leg 3–0 as Stevie Chalmers scored and Gemmell got a brace. Celtic faced French side Nantes in the second round, and won the away leg 3–1. Nantes had taken the lead through Francis Magny, before McBride had equalised to level the match at 1–1. In the second half, Bobby Lennox and Bertie Auld scored to seal a victory. Celtic won the home leg by the same scoreline. Jimmy Johnstone put them in front, before Gérard Georgin equalised. Celtic again scored twice in the second half, as Chalmers and Lennox secured the victory.[2]
Celtic faced Yugoslavia (now Serbian) side Vojvodina Novi Sad in the quarter-finals, and lost the first leg 1–0 after a goal from Milan Stanić; this was Celtic's only defeat of the competition. The tie looked like it was going to end in a draw after Chalmers had given Celtic a 1–0 lead in the second leg. This would have resulted in the teams from having to go to Rotterdam for a replay. However, in the 90th minute captain Billy McNeill scored to give Celtic the victory.[3] In the semi-finals, Czechoslovakian side Dukla Prague were beaten 3–1 in Glasgow, Johnstone put the hosts in front, before Stanislav Štrunc equalised. A second half brace from Willie Wallace gave Celtic the victory. The teams then drew 0–0 in Prague, which meant Celtic progressed to the final.[2][4]
Internazionale[edit]
Round
Opponents
First leg
Second leg
Aggregate score
First round
Torpedo Moscow 1–0 (h) 0–0 (a) 1–0
Second round
Vasas 2–1 (h) 2–0 (a) 4–1
Quarter-finals
Real Madrid 1–0 (h) 2–0 (a) 3–0
Semi-finals
CSKA Red Flag 1–1 (a) 1–1 (h) 2–2
Play-off
CSKA Red Flag 1–0 (n)
Internazionale had won the 1965–66 Serie A, their tenth title, by four points over second placed Bologna.[5] As a result of this, they qualified for the European Cup and their first round opponents were Soviet side Torpedo Moscow. Internazionale won the first leg 1–0, thanks to an own goal by Valery Voronin before drawing 0–0 in Russia. Their second round opponents were Vasas of Hungary, Internazionale won 2–1 at home, with goals from Carlo Soldo and Mario Corso, while Lajos Puskás had scored for the visitors. Two goals from Sandro Mazzola gave Inter the victory in the second leg.
Internazionale beat six-time champions and holders Real Madrid in the quarter-finals. Internazionale won 1–0 at home, through a Renato Cappellini goal. Before defeating Madrid 2–0 in Spain, thanks to another goal from Cappellini, and an own goal from Ignacio Zoco. In the semi-finals. Internazionale faced Bulgarian side CSKA Red Flag (now CSKA Sofia). Giacinto Facchetti scored for Internazionale, as they drew 1–1 at home, with Nikola Tsanev scoring for the visitors. Facchetti scored again in Bulgaria, but his goal was cancelled out by Nikolay Radlev, meaning that a play-off was needed to settle the tie. The play-off was supposed to be held in Graz, Austria, but CSKA were persuaded to let it be moved to Bologna in Italy, after they were offered a larger share of the gate money.[6] The match was won 1–0 by Internazionale, thanks to a goal from Cappellini, sealing their place in the final.[7]
Match[edit]
Background[edit]
The final was held in the Estádio Nacional, Lisbon
Internazionale had won the European Cup in two of the previous three seasons, 1964 and 1965. Pre-match talk focused on Internazionale winning a famous tripletta of European Cups and they were considered strong favourites going into the game.[8][9]
Internazionale were very well known for using a defensive tactic, the Catenaccio, which meant that they won many matches by slim scorelines and rarely conceded.[10] Their manager, Helenio Herrera, was the highest paid in Europe and was considered to be the catalyst of their success.[11] By contrast Celtic were an attacking team. Before the match their manager Jock Stein said that, "Celtic will be the first team to bring the European Cup back to Britain... we are going to attack as we have never attacked before,"[10]
One of Celtic's most important players, striker Joe McBride was to miss the match. He had suffered a long-term knee injury and his last match of the season was on 24 December 1966. McBride would definitely have played had he been fit. He was in such good form for Celtic that, despite missing half the season, McBride finished as the top scorer in Scotland that year with 35 goals in 26 appearances.[12]
Both Internazionale and Celtic had been performing well domestically throughout the season. Only a few days before the final Internazionale had been on the verge of winning a historic treble but losses in their last two games knocked them out of the semi-finals of the Coppa Italia and cost them the Scudetto. The European Cup was the last chance for them to redeem what had initially been such a promising season. Celtic came into the final having already won the First Division, the Scottish Cup and the Scottish League Cup as well as the Glasgow Cup earlier in the season.[13][14]
Summary[edit]
Internazionale had the first attack of the match. With Renato Cappellini running down the wing and supplying a cross to Sandro Mazzola whose header hit Celtic goalkeeper Ronnie Simpson's knees. Internazionale won a penalty moments later. Jim Craig fouled Cappellini in the box, and Mazzola converted to put Inter in front after only six minutes. Once they had taken the lead, Internazionale retreated back into their defensive style, which allowed Celtic to attack. However, they struggled to get through Internazionale's defensive wall and were mainly restricted to long shots from outside the box. Bertie Auld hit the crossbar then a cross from Jimmy Johnstone was gathered up by Giuliano Sarti, who then tipped a header from the same player over the crossbar. Internazionale pulled nine men back, but Celtic kept attacking them. Tommy Gemmell's dangerous free-kick was saved by Sarti, he then speculatively attempted to lob the Inter 'keeper and hit the bar. Despite Celtic's inability to break through their opponents defence they were in complete control of the match and Internazionale were not able to attack. Inter had not had another chance since their goal, whilst Celtic found themselves foiled time and again by outstanding goalkeeping from Sarti.
After just over an hour, Gemmell finally managed to equalise for Celtic when Craig passed to him from the right wing and he scored with a powerful 25 yard shot.[15] The balance of play remained the same with Inter defending deeply against sustained Celtic attacking. With about five minutes remaining, a long-range shot from Bobby Murdoch was diverted by Stevie Chalmers past a wrong-footed Sarti. This proved to be the winning goal.[16]
Details[edit]
Celtic v Internazionale
25 May 1967
17:30 WEST
Celtic
2–1
Internazionale
Gemmell 63'
Chalmers 84' Report Mazzola 7' (pen.)
Estádio Nacional, Lisbon
Attendance: 45,000
Referee: Kurt Tschenscher (West Germany)
Celtic
Internazionale
GK 1 Ronnie Simpson
RB 2 Jim Craig
LB 3 Tommy Gemmell
CM 4 Bobby Murdoch
CB 5 Billy McNeill (c)
CB 6 John Clark
RF 7 Jimmy Johnstone
CF 8 Willie Wallace
CF 9 Stevie Chalmers
CM 10 Bertie Auld
LF 11 Bobby Lennox
Manager:
Jock Stein
GK 1 Giuliano Sarti
RB 2 Tarcisio Burgnich
LB 3 Giacinto Facchetti
CM 4 Gianfranco Bedin
CB 5 Aristide Guarneri
SW 6 Armando Picchi (c)
RF 7 Angelo Domenghini
CM 8 Sandro Mazzola
CF 9 Renato Cappellini
CM 10 Mauro Bicicli
LF 11 Mario Corso
Manager:
Helenio Herrera
Assistant referees:
Rudibert Jacobi (West Germany)
Rudolf Eisemann (West Germany)
Post-match[edit]
After the final whistle, there was a pitch invasion by Celtic fans, which meant that the Celtic team could not be presented the trophy on the pitch.[17] Some of the Celtic players also had their shirts taken by Celtic supporters.[17] Celtic captain Billy McNeill had to be ushered around the outside of the stadium under armed guards to receive the trophy on a podium in the stand.[17]
Internazionale's loss in the final is considered to be the downfall of "La Grande Inter", the greatest period of success in the club's history. They had been one of the top teams in Europe for the previous three years, however, failed to recover from their bad season in which they lost out to Celtic as well as in their two domestic competitions.[18] They finished the following season trophyless again and Helenio Herrera, the manager who was considered to be the catalyst of their success, then left the club.[13]
Celtic's attacking style play against Internazionale's catenaccio was heralded as a win for football. Internazionale manager Helenio Herrera said that "We can have no complaints. Celtic deserved their victory. We were beaten by Celtic's force. Although we lost, the match was a victory for sport."[13] while one Portuguese official said "This attacking play, this is the real meaning of football. This is the true game.". The Portuguese newspaper, Mundo Desportivo, said "It was inevitable. Sooner or later the Inter of Herrera, the Inter of catenaccio, of negative football, of marginal victories, had to pay for their refusal to play entertaining football."[19]
Celtic manager Jock Stein received widespread praise following the final. Liverpool manager Bill Shankly said to him after the match, "John, you're immortal now". Since the match a stand has been named after him at Celtic Park. While Stein also received the Commander of the Order of the British Empire.[20] Stein is considered by many, including Alex Ferguson, to be the greatest ever Scottish manager, with his victory in the final being one of the main reasons for this.[21][22]
The Celtic team from that year has also received much recognition. They have become known as the Lisbon Lions and are widely considered the greatest team in Celtic's history. All of Celtic's players were born within a 30-mile radius of Glasgow.[23] In 2000, Celtic named a stand at Celtic Park after the Lisbon Lions.[24] They also won the BBC Sports Personality Team of the Year Award in 1967.[25]
See also[edit]
1966–67 European Cup
1967 European Cup Winners' Cup Final
1967 Inter-Cities Fairs Cup Final
References[edit]
Jump up ^ "Scotland 1965/66". Rec.Sport.Soccer Statistics Foundation. Retrieved 27 September 2011.
^ Jump up to: a b "Champions Cup 1966–67". Rec.Sport.Soccer Statistics Foundation. Retrieved 27 September 2011.
Jump up ^ "Glasgow's record breakers". Glasgow Evening Times. 7 May 2007. Retrieved 28 September 2011.
Jump up ^ "1966/67 European Champions Clubs' Cup". UEFA.com. Retrieved 27 September 2011.
Jump up ^ "Italy 1965/66". Rec.Sport.Soccer Statistics Foundation. Archived from the original on 21 September 2011. Retrieved 27 September 2011.
Jump up ^ "Season 1966–67". European Cup History. Retrieved 28 November 2011.
Jump up ^ "1966/67 European Champions Clubs' Cup". UEFA.com. Retrieved 27 September 2011.
Jump up ^ "Lions' legend lives on". FIFA.com. 17 July 2007. Retrieved 2 December 2011.
Jump up ^ Glanville, Brian (17 May 2001). "Obituary: Bobby Murdoch". London: The Guardian. Retrieved 13 November 2011.
^ Jump up to: a b "LOCAL HEROES: THE LISBON LIONS". Chris Hunt. June 2007. Retrieved 13 November 2011.
Jump up ^ Fox, Norman (11 November 1997). "Obituary: Helenio Herrera". The Independent. London. Retrieved 2 December 2011.
Jump up ^ "Missing out on being Lisbon Lion saved my life, says Celtic legend Joe McBride". Daily Record. 23 May 2010. Retrieved 28 November 2011.
^ Jump up to: a b c "Jock Stein's Lisbon Lions shock Inter". ESPN. 19 May 2010. Retrieved 2 December 2011.
Jump up ^ "Top ten successful seasons". Sky Sports. 26 March 2009. Retrieved 10 April 2012.
Jump up ^ Barham, Albert (25 May 1967). "Relentless attack captures European Cup". London: The Guardian. Retrieved 28 November 2011.
Jump up ^ Forsyth, Roddy (15 May 2001). "Murdoch the true Lionheart". London: The Telegraph. Retrieved 7 April 2012.
^ Jump up to: a b c "Celtic win European Cup 1967". BBC Sport. Retrieved 10 February 2013.
Jump up ^ Rinaldi, Giancarlo (23 May 2007). "Inter scarred by Celtic memories". BBC Sport. British Broadcasting Corporation. Retrieved 2 December 2011.
Jump up ^ "From The Vault: The Lisbon Lions". London: The Guardian. 21 May 2009. Retrieved 2 December 2011.
Jump up ^ "March 9 – Jock Stein Takes Over Celtic". On this football day. 9 March 2009. Retrieved 2 December 2011.
Jump up ^ "I'll never be as good as Big Jock Says SIR ALEX FERGUSON; MAN UTD v SPURS Wembley, kick-off 3pm Live on Sky Sports CARLING CUP". Sunday Mail. 1 March 2009. Retrieved 10 April 2012.
Jump up ^ "1967 European Cup win makes Jock Stein greatest Scots manager of all time, says Lou Macari". Daily Record. 5 March 2011. Retrieved 10 April 2012.
Jump up ^ "1967: Celtic win European Cup". On This Day. British Broadcasting Corporation. 25 May 2005. Retrieved 6 December 2011.
Jump up ^ Reynolds, Jim (28 January 2007). "Fans will now roar from the Lisbon Lions Stand". The Herald. Retrieved 6 April 2012.
Jump up ^ "The Lisbon Lions". Daily Mirror. Retrieved 2 December 2011.
[show]
v
t
e
1966–67 in European football (UEFA)
« 1965–66
1967–68 »
Domestic leagues
Albania
Austria
Belgium
Bulgaria
Cyprus
Czechoslovakia
Denmark '66 '67
England
Faroe Islands '66 '67
Finland '66 '67
France
East Germany
West Germany
Greece
Hungary '66 '67
Iceland '66 '67
Israel
Italy
Luxembourg
Malta
Netherlands
Northern Ireland
Norway '66 '67
Poland
Portugal
Republic of Ireland
Romania
Scotland
Soviet Union '66 '67
Spain
Sweden '66 '67
Switzerland
Turkey
Yugoslavia
Domestic cups
Austria
Belgium
Bulgaria
Cyprus
Czechoslovakia
Denmark
England
Faroe Islands '66 '67
Finland '66 '67
France
East Germany
West Germany
Greece
Hungary '66 '67
Iceland '66 '67
Israel
Italy
Liechtenstein
Luxembourg
Malta
Netherlands
Northern Ireland
Norway '66 '67
Poland
Portugal
Republic of Ireland
Romania
San Marino
Scotland
Soviet Union
Spain
Sweden '67
Switzerland
Turkey
Wales
Yugoslavia
League cups
England
Scotland
UEFA competitions
European Cup (Final)
Cup Winners' Cup (Final)
Non-UEFA competitions
Inter-Cities Fairs Cup (Final)
Balkans Cup
Mitropa Cup
[show]
v
t
e
European Cup and UEFA Champions League
European Cup era, 1955–1992
Seasons
1955–56
1956–57
1957–58
1958–59
1959–60
1960–61
1961–62
1962–63
1963–64
1964–65
1965–66
1966–67
1967–68
1968–69
1969–70
1970–71
1971–72
1972–73
1973–74
1974–75
1975–76
1976–77
1977–78
1978–79
1979–80
1980–81
1981–82
1982–83
1983–84
1984–85
1985–86
1986–87
1987–88
1988–89
1989–90
1990–91
1991–92
Finals
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
UEFA Champions League era, 1992–present
Seasons
1992–93
1993–94
1994–95
1995–96
1996–97
1997–98
1998–99
1999–2000
2000–01
2001–02
2002–03
2003–04
2004–05
2005–06
2006–07
2007–08
2008–09
2009–10
2010–11
2011–12
2012–13
2013–14
2014–15
2015–16
2016–17
2017–18
2018–19
Finals
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
2016
2017
2018
2019
2020
History
Finals
Winning managers
Winning players
Records and statistics
Top scorers
Hat-tricks
Performance comparison
Anthem
Broadcasters
Trophy
[show]
v
t
e
International men's club football
FIFA
FIFA Club World Cup
List of association football clubs
Africa
CAF – Champions League
Confederation Cup
Super Cup
Top-division clubs
Asia
AFC – Champions League
AFC Cup
Top-division clubs
Europe
UEFA – Champions League
Europa League
Super Cup
Top-division clubs
North,
Central America
and the Caribbean
CONCACAF – Champions League
CONCACAF League
Top-division clubs
Oceania
OFC – Champions League
Top-division clubs
South America
CONMEBOL – Copa Libertadores
Copa Sudamericana
Recopa Sudamericana
Top-division clubs
See also: International women's club football
[show]
v
t
e
F.C. Internazionale Milano matches
Italian football championship Final
1920
Coppa Italia Finals
1939
1959
1965
1977
1978
1982
2000
2005
2006
2007
2008
2010
2011
Supercoppa Italiana
1989
2000
2005
2006
2007
2008
2009
2010
2011
UEFA Champions League Finals
1964
1965
1967
1972
2010
UEFA Cup Finals
1991
1994
1997
1998
UEFA Super Cup
2010
Intercontinental Cups
1964
1965
Intercontinental Supercup
1968
FIFA Club World Cup Final
2010
[show]
v
t
e
Celtic F.C. matches
Scottish Cup Finals
1889
1892
1893
1894
1899
1900
1901
1902
1904
1907
1908
1909
1911
1912
1914
1923
1925
1926
1927
1928
1931
1933
1937
1951
1954
1955
1956
1961
1963
1965
1966
1967
1969
1970
1971
1972
1973
1974
1975
1977
1980
1984
1985
1988
1989
1990
1995
1999
2001
2002
2004
2005
2007
2011
2013
2017
Scottish League Cup Finals
1956
1957
1964
1965
1966
1967
1969 (April)
1969 (October)
1970
1971
1972
1973
1974
1975
1976
1978
1982
1984 (March)
1986
1990
1994
1997
2000
2001
2003
2006
2009
2011
2012
2015
2016 (November)
2017
European Cup Finals
1967
1970
UEFA Cup Final
2003
Intercontinental Cup
1967
Other matches
Coronation Cup Final (1953)
CNE Cup of Champions (1968)
Rangers 2–2 Celtic (1987)
2007 MLS All-Star Game
Lincoln Red Imps 1–0 Celtic (2016)
Retrieved from "https://en.wikipedia.org/w/index.php?title=1967_European_Cup_Final&oldid=807992669"
Categories:
1966–67 in European football
Celtic F.C. matches
International club association football competitions hosted by Portugal
Inter Milan matches
UEFA Champions League Finals
1966–67 in Portuguese football
1966–67 in Scottish football
1966–67 in Italian football
May 1967 sports events
Sports competitions in Lisbon
1960s in Lisbon
Hidden categories:
Good articles
Navigation menu
Personal tools
Not logged in
Talk
Contributions
Create account
Log in
Namespaces
Article
Talk
Variants
Views
Read
Edit
View history
More
Search
Navigation
Main page
Contents
Featured content
Current events
Random article
Donate to Wikipedia
Wikipedia store
Interaction
Help
About Wikipedia
Community portal
Recent changes
Contact page
Tools
What links here
Related changes
Upload file
Special pages
Permanent link
Page information
Wikidata item
Cite this page
Print/export
Create a book
Download as PDF
Printable version
Languages
العربية
Беларуская (тарашкевіца)
Dansk
Español
فارسی
Français
Հայերեն
हिन्दी
Hrvatski
Bahasa Indonesia
Italiano
עברית
Magyar
Nederlands
Português
Română
Русский
Türkçe
Tiếng Việt
中文
Edit links
This page was last edited on 31 October 2017, at 06:49.
Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation, Inc., a non-profit organization.
Privacy policy
About Wikipedia
Disclaimers
Contact Wikipedia
Developers
Cookie statement
Mobile view
Enable previews

values: mw-page-base noprint

values: mw-head-base noprint

values: This is a good article. Follow the link for more information.

attrs: class id

values: firstHeading firstHeading en

values: siteSub noprint

values: jump-to-nav mw-jump

value: caption

attrs: href class

value: td

attrs: colspan style

value: tr

value: th

attrs: scope style

values:
row
white-space: nowrap

attrs: width style

attrs: href class title

values:
/wiki/F.C._Internazionale_Milano
mw-redirect
F.C. Internazionale Milano

values:
background-color: inherit
100%
nowraplinks

values:
/wiki/Brace_(sports)
mw-redirect
Brace (sports)

values:
/wiki/William_Wallace_(footballer)
mw-redirect
William Wallace (footballer)

values:
width:39%; text-align:right
vcard attendee

values:
width:39%; text-align:left
vcard attendee

values: float:left; width:61%; table-layout:fixed; text-align:center

values: width: 100%; background-color: transparent; clear:both

attrs: alt src title width height style data-file-width data-file-height

values: position: absolute; left: 0px; top: 0px; width: 31px; height: 59px; background-color: #FFFFFF;

attrs: alt src width height style srcset data-file-width data-file-height

attrs: alt src width height style data-file-width data-file-height

values: position: absolute; left: 31px; top: 0px; width: 38px; height: 59px; background-color: #00A200;

values: position: absolute; left: 69px; top: 0px; width: 31px; height: 59px; background-color: #FFFFFF;

values: position: absolute; left: 0px; top: 59px; width: 100px; height: 36px; background-color: #FFFFFF

values: position: absolute; left: 0px; top: 95px; width: 100px; height: 40px; background-color: #FFFFFF

values: position: relative; left: 0px; top: 0px; width: 100px; height: 135px; margin: 0 auto; padding: 0;

values: width: 100px; margin: 0 auto; padding: 0;

values: position: absolute; left: 0px; top: 0px; width: 31px; height: 59px; background-color: #3344CC;

values: position: absolute; left: 31px; top: 0px; width: 38px; height: 59px; background-color: #3344CC;

values: position: absolute; left: 69px; top: 0px; width: 31px; height: 59px; background-color: #3344CC;

values: position: absolute; left: 0px; top: 59px; width: 100px; height: 36px; background-color: #000000

values: position: absolute; left: 0px; top: 95px; width: 100px; height: 40px; background-color: #3344CC

values: BBC Sports Personality Team of the Year Award

values: -moz-column-width: 30em; -webkit-column-width: 30em; column-width: 30em; list-style-type: decimal;

values: Template:1966–67 in European football (UEFA)

values: Template talk:1966–67 in European football (UEFA)

values: Template:1965–66 in European football (UEFA)

values: Template:1967–68 in European football (UEFA)

values: 1966–67 Austrian Cup (page does not exist)

values: 1966–67 Belgian Cup (page does not exist)

values: 1966–67 Czechoslovak Cup (page does not exist)

values: 1966–67 Danish Cup (page does not exist)

values: 1966 Faroe Islands Cup (page does not exist)

values: 1967 Faroe Islands Cup (page does not exist)

values: 1966–67 FDGB-Pokal (page does not exist)

values: 1966 Icelandic Cup (page does not exist)

values: 1967 Icelandic Cup (page does not exist)

values: 1966–67 Coppa Italia (page does not exist)

values: 1966–67 Liechtenstein Cup (page does not exist)

values: 1966–67 Luxembourg Cup (page does not exist)

values: 1966–67 Maltese FA Trophy (page does not exist)

values: 1966–67 Polish Cup (page does not exist)

values: 1966–67 Coppa Titano (page does not exist)

values: 1966–67 Soviet Cup (page does not exist)

values: 1966–67 Turkish Cup (page does not exist)

values: nowraplinks collapsible autocollapse navbox-inner

values:
collapseButton1
#
color: rgb(240, 240, 240);

values: Template talk:UEFA Champions League seasons

values: European Cup and UEFA Champions League history

values: List of European Cup and UEFA Champions League finals

values: List of European Cup and UEFA Champions League winning managers

values: List of European Cup and UEFA Champions League winning players

values: European Cup and UEFA Champions League records and statistics

values: List of European Cup and UEFA Champions League top scorers

values: List of UEFA Champions League hat-tricks

values: UEFA Champions League clubs performance comparison

values: List of UEFA Champions League broadcasters

values: nowraplinks hlist collapsible autocollapse navbox-inner

values: Template talk:International club football

values: width:1%;line-height: 1em;-moz-box-shadow: inset -5px 0 0 0 #deb887; -webkit-box-shadow: inset -5px 0 0 0 #deb887; box-shadow: inset -5px 0 0 0 #deb887;

values: List of top-division football clubs in CAF countries

values:
navbox-image
6
width:1px;padding:0px 0px 0px 2px

values: width:1%;line-height: 1em;-moz-box-shadow: inset -5px 0 0 0 #ffb6c1; -webkit-box-shadow: inset -5px 0 0 0 #ffb6c1; box-shadow: inset -5px 0 0 0 #ffb6c1;

values: List of top-division football clubs in Asian Football Confederation members

values: width:1%;line-height: 1em;-moz-box-shadow: inset -5px 0 0 0 #4682b4; -webkit-box-shadow: inset -5px 0 0 0 #4682b4; box-shadow: inset -5px 0 0 0 #4682b4;

values: List of top-division football clubs in UEFA countries

values: width:1%;line-height: 1em;-moz-box-shadow: inset -5px 0 0 0 #db7093; -webkit-box-shadow: inset -5px 0 0 0 #db7093; box-shadow: inset -5px 0 0 0 #db7093;

values: List of top-division football clubs in CONCACAF countries

values: width:1%;line-height: 1em;-moz-box-shadow: inset -5px 0 0 0 #ffd700; -webkit-box-shadow: inset -5px 0 0 0 #ffd700; box-shadow: inset -5px 0 0 0 #ffd700;

values: List of top-division football clubs in OFC countries

values: width:1%;line-height: 1em;-moz-box-shadow: inset -5px 0 0 0 #8fbc8f; -webkit-box-shadow: inset -5px 0 0 0 #8fbc8f; box-shadow: inset -5px 0 0 0 #8fbc8f;

values: List of top-division football clubs in CONMEBOL countries

values: Template:International women's club football

values:
collapseButton3
#
color: rgb(255, 255, 255);

values: Template:F.C. Internazionale Milano matches

values: Template talk:F.C. Internazionale Milano matches

values: 1939 Coppa Italia Final (page does not exist)

values: 1959 Coppa Italia Final (page does not exist)

values: 1965 Coppa Italia Final (page does not exist)

values: 1977 Coppa Italia Final (page does not exist)

values:
/wiki/1978_Coppa_Italia_Final
mw-redirect
1978 Coppa Italia Final

values: 1982 Coppa Italia Final (page does not exist)

values:
/wiki/Intercontinental_Supercup
mw-redirect
Intercontinental Supercup

values: background:#008000; color: white;border:1px solid white;;;background:none transparent;border:none;-moz-box-shadow:none;-webkit-box-shadow:none;box-shadow:none;

values: background:#008000; color: white;border:1px solid white;;width:1%

values: 1969 Scottish League Cup Final (October)

values: 2016 Scottish League Cup Final (November)

values: Category:International club association football competitions hosted by Portugal

values: Discussion about edits from this IP address [ctrl-alt-n]

values: A list of edits made from this IP address [ctrl-alt-y]

values: You are encouraged to create an account and log in; however, it is not mandatory

values: You're encouraged to log in; however, it's not mandatory. [ctrl-alt-o]

values:
/wiki/1967_European_Cup_Final
View the content page [ctrl-alt-c]
c

values: Discussion about the content page [ctrl-alt-t]

values: p-namespaces navigation vectorTabs p-namespaces-label

values:
p-variants
navigation
vectorMenu emptyPortlet
p-variants-label

values: Past revisions of this page [ctrl-alt-h]

values: p-views navigation vectorTabs p-views-label

values:
p-cactions
navigation
vectorMenu emptyPortlet
p-cactions-label

values:
mw-wiki-logo
/wiki/Main_Page
Visit the main page

values:
/wiki/Main_Page
Visit the main page [ctrl-alt-z]
z

values: Featured content – the best of Wikipedia

values: Find background information on current events

values:
/wiki/Special:Random
Load a random article [ctrl-alt-x]
x

values: portal navigation p-navigation p-navigation-label

values: Guidance on how to use and edit Wikipedia

values: About the project, what you can do, where to find things

values: A list of recent changes in the wiki [ctrl-alt-r]

values: portal navigation p-interaction p-interaction-label

values: List of all English Wikipedia pages containing links to this page [ctrl-alt-j]

values: Recent changes in pages linked from this page [ctrl-alt-k]

values:
/wiki/Wikipedia:File_Upload_Wizard
Upload files [ctrl-alt-u]
u

values: A list of all special pages [ctrl-alt-q]

values: Permanent link to this revision of the page

values: Link to connected data repository item [ctrl-alt-g]

values: Printable version of this page [ctrl-alt-p]

values:
uls-settings-trigger
Language settings
0
button
true

values: نهائي كأس أوروبا 1967 – Arabic

values: Фінал Кубка эўрапейскіх чэмпіёнаў 1967 году – Belarusian (Taraškievica orthography)

values: Mesterholdenes Europa Cup finale 1967 – Danish

values: Final de la Copa de Campeones de Europa 1966-67 – Spanish

values: interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle

values: فینال جام باشگاه‌های اروپا ۱۹۶۷ – Persian

values: Finale de la Coupe des clubs champions européens 1966-1967 – French

values: Եվրոպայի չեմպիոնների գավաթի եզրափակիչ 1967 – Armenian

values: १९६७ यूरोपीय कप फाइनल – Hindi

values: Final Piala Champions Eropa 1967 – Indonesian

values: Finale della Coppa dei Campioni 1966-1967 – Italian

values: גמר גביע אירופה לאלופות 1967 – Hebrew

values: 1967-es bajnokcsapatok Európa-kupája-döntő – Hungarian

values: Final da Taça dos Clubes Campeões Europeus de 1966-67 – Portuguese

values: Finala Cupei Campionilor Europeni 1967 – Romanian

values: Финал Кубка европейских чемпионов 1967 – Russian

values: 1967 Şampiyon Kulüpler Kupası Finali – Turkish

values: Chung kết Cúp C1 châu Âu 1967 – Vietnamese

values: mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject page-1967_European_Cup_Final rootpage-1967_European_Cup_Final skin-vector action-view