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Functions downstream of the tRNA-splicing endonuclease that removes introns, ligating the two generated halves via phosphodiester bond formation. The ligation reaction, which requires guanosine triphosphate (GTP) and Mn(2+), proceeds through three metal-dependent steps. The first step requires ZBTB8OS/Archease and involves the guanylylation of RTCB at its active site histidine, forming a covalent GMP-histidine intermediate. Before the second step, RTCB also hydrolyzes the 2',3'-cyclic phosphate (cP) at the 3' end of the 5' tRNA exon, typically generated by the tRNA-splicing endonuclease, producing a 3' phosphate. The covalently bound GMP is then transferred to this 3' phosphate to form an RNA(3')-P-P-(5')G intermediate. In the final step, the 5'-OH of the 3' exon attacks the activated 3' end of the 5' exon, forming a 3'-5' phosphodiester bond and releasing GMP. RTCB also functions in non-canonical, spliceosome-independent, cytoplasmic splicing of XBP1 mRNAs during the unfolded protein response (UPR). Upon endoplasmic reticulum (ER) stress, the endoribonuclease IRE1/ERN1 excises a short intron, generating free exon ends that are aligned by RNA-intrinsic, zipper-like stem-loop structures. These exon ends are then recognized and ligated by RTCB. This splicing event yields the active XBP1s transcription factor, which induces genes required to resolve protein folding defects in the endoplasmic reticulum"}],"commentType":"FUNCTION"},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"a 3'-end 3'-phospho-ribonucleotide-RNA + a 5'-end dephospho-ribonucleoside-RNA + GTP = a ribonucleotidyl-ribonucleotide-RNA + GMP + 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2 Mn(2+) per subunit. Three Mn(2+) ions are bound in the composite active site formed by RTCB and Archease. Two are exclusively coordinated by RTCB residues and the third one is solely coordinated by ZBTB8OS/Archease residues."}]}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q9Y3I0"}],"value":"ZBTB8OS/Archease stimulates the activity of the tRNA ligase complex by participating to RTCB active site guanylylation reaction. While Mg(2+), Ca(2+) and to a lesser extent Co(2+) show low levels of covalent incorporation of GMP the strongest guanylylation occurs in presence of Mn(2+) ions"}],"commentType":"ACTIVITY REGULATION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q9Y3I0"}],"value":"Catalytic component of the tRNA-splicing ligase core complex composed of RTCB and the accessory proteins DDX1, C2orf49/Ashwin, FAM98B and RTRAF/CGI-99. Interacts with PYROXD1; the interaction is direct and shields Cys-122 preventing oxidative inactivation of the catalytic center. The interaction could also regulate RTCB activation and depends on divalent cations. Interacts with ZBTB8OS/Archease; active site guanylylation reaction relies on the activation factor ZBTB8OS/Archease"}],"commentType":"SUBUNIT"},{"commentType":"SUBCELLULAR LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q9Y3I0"}],"value":"Enters into the nucleus in case of active transcription while it accumulates in cytosol when transcription level is low"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q9Y3I0"}],"value":"Nucleus","id":"SL-0191"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18508721"}],"value":"Cytoplasm","id":"SL-0086"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21046479"}],"value":"Ubiquitously expressed. Highly expressed in the epididymis with predominant enrichment in the initial segment. During sexual maturation, it is expressed in the caput epididymides"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21046479"}],"value":"Down-regulated in the epididymis upon castration"}],"commentType":"INDUCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20510672"}],"value":"Essential during post-implantation development of embryos"}],"commentType":"DISRUPTION PHENOTYPE"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03144"}],"value":"Belongs to the RtcB family"}],"commentType":"SIMILARITY"}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":505,"modifier":"EXACT"}},"description":"RNA-splicing ligase RTCB","featureId":"PRO_0000255243"},{"type":"Active site","location":{"start":{"value":428,"modifier":"EXACT"},"end":{"value":428,"modifier":"EXACT"}},"description":"GMP-histidine 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