description
Human ribosomal RNAs (rRNAs) contain about 200 residues that are enzymatically modified after transcription in the nucleolus (Maden and Khan 1977, Maden 1988, Maden and Hughes 1997, reviewed in Hernandez-Verdun et al. 2010, Boschi-Muller and Motorin 2013). The modified residues occur in regions of the rRNAs that are located in functionally important parts of the ribosome, notably in the A and P peptidyl transfer sites, the polypeptide exit tunnel, and intersubunit contacts (Polikanov et al. 2015, reviewed in Decatur and Fournier 2002, Chow et al. 2007, Sharma and Lafontaine 2015). The two most common modifications are pseudouridines and 2'-O-methylribonucleotides. Formation of pseudouridine from encoded uridine is catalyzed by box H/ACA small nucleolar ribonucleoprotein (snoRNP) complexes (reviewed in Hamma and Ferre-D'Amare 2010, Watkins and Bohnsack 2011, Ge and Yu 2013, Kierzek et al. 2014, Yu and Meier 2014) and methylation of the hydroxyl group of the 2' carbon is catalyzed by box C/D snoRNPs (Kiss-Laszlo et al. 1996, Lapinaite et al. 2013, reviewed in Watkins and Bohnsack 2011). The snoRNP complexes contain common sets of protein subunits and unique snoRNAs that guide each complex to its target nucleotide of the rRNA by base-pairing between the snoRNA and the rRNA (reviewed in Henras et al. 2004, Watkins and Bohnsack 2011). Other modifications of rRNA include 5-methylcytidine (reviewed in Squires and Preiss 2010), 1-methylpseudouridine, 7-methylguanosine, 6-dimethyladenosine, and 4-acetylcytidine (reviewed in Sharma and Lafontaine 2015). In yeast most modifications are introduced co-transcriptionally (Kos and Tollervey 2010, reviewed in Turowski and Tollervey 2015), however the order of modification events and pre-rRNA cleavage events is not well characterized

external resources
NCBI:1383087
REACTOME:R-HSA-6790901
PUBMED:25590339
PUBMED:15193314
PUBMED:20347423
PUBMED:25775268
PUBMED:21956940
PUBMED:24490730
PUBMED:19917616
PUBMED:25176256
PUBMED:3418702
PUBMED:26410597
PUBMED:22122054
PUBMED:23391857
PUBMED:8674114
PUBMED:12114023
PUBMED:17894445
PUBMED:24121435
PUBMED:412496
PUBMED:9211966
PUBMED:22065625
PUBMED:24369424

genes
DKC1 , FBL , SNU13 , NOP2 , PWP2 , RPS2 , RPS6 , RPS7 , RPS9 , RPS14 , NOP14 , RRP9 , WDR46 , UTP14C , BMS1 , RCL1 , MPHOSPH10 , EMG1 , NOP56 , TBL3 , UTP14A , WDR3 , DDX52 , KRR1 , PDCD11 , WDR43 , DCAF13 , NOL11 , UTP25 , DIMT1 , UTP20 , RRP7A , FCF1 , UTP18 , UTP11 , DDX47 , TRMT112 , NOP58 , GAR1 , DDX49 , HEATR1 , NAT10 , IMP3 , NOP10 , THUMPD1 , NHP2 , UTP6 , UTP3 , DHX37 , NOL6 , NOC4L , WDR75 , UTP15 , UTP4 , RRP36 , IMP4 , BUD23 , TSR3 , WDR36 , MIR1292 , MIR664B ,