Standard name Systematic name Name description Description
TUF1 YOR187W Mitochondrial translation elongation factor Tu (EF-Tu) Mitochondrial translation elongation factor Tu (EF-Tu); involved in fundamental pathway of mtDNA homeostasis; comprises both GTPase and guanine nucleotide exchange factor activities, while these activities are found in separate proteins in S. pombe and humans; rare mutations in human mitochondrial elongation factor Tu (EFTu) associated with severe lactic acidosis, rapidly progressive fatal encephalopathy, severe infantile macrocystic leukodystrophy with micropolygyria
Visualize the interactome of TUF1
Links to external databases:
SGD   |   KEGG   |  

Cell cycle phase and timing of transcription peak

Cell cycle phase Time (min)
G2 108

CYCLoPs localization & abundance

WT1 WT2 WT3
Golgi 0.15 NaN NaN
Mito 0.95 NaN 0.880
Nucleolus 0.15 NaN NaN
Vac/Vac Memb 0.15 NaN 0.064
Nuc Periphery NaN NaN 0.080

GFP localization & abundance

GFP localization GFP abundance
mitochondrion 58500

GO term count

Metabolism Cell cycle Cell division Signal transduction DNA replication
TUF1 4 0 0 0 0

List of interactors

Use the search utility to find the interactor you are looking for. By clicking the column headers the table will be sorted on that column. Use shift+click to sort on multiple columns. Default sorting is on number of experiments, number of unique methods and the number of publications, and alphabetical on standard name, in that order.
Standard name Systematic name Name description Type #Experiments #Methods #Publications Evidence
HSP82 YPL240C Hsp90 chaperone physical 4 1 4 Affinity Capture-MS, Affinity Capture-MS, Affinity Capture-MS, Affinity Capture-MS
PUF3 YLL013C Protein of the mitochondrial outer surface physical 4 1 4 Affinity Capture-RNA, Affinity Capture-RNA, Affinity Capture-RNA, Affinity Capture-RNA
FKH1 YIL131C Forkhead family transcription factor regulation 3 2 3 chromatin immunoprecipitation-chip evidence, chromatin immunoprecipitation-chip evidence, ChEC-Seq experiments
DSK2 YMR276W Nuclear-enriched ubiquitin-like polyubiquitin-binding protein genetic 2 2 2 Positive Genetic, Negative Genetic
RPN11 YFR004W Metalloprotease subunit of 19S regulatory particle physical 2 1 2 Affinity Capture-MS, Affinity Capture-MS
SSA1 YAL005C ATPase involved in protein folding and NLS-directed nuclear transport physical 2 1 2 Affinity Capture-MS, Affinity Capture-MS
ATP2 YJR121W Beta subunit of the F1 sector of mitochondrial F1F0 ATP synthase genetic 2 1 1 Synthetic Rescue, Synthetic Rescue
TAF9 YMR236W Subunit (17 kDa) of TFIID and SAGA complexes genetic 2 1 1 Synthetic Lethality, Synthetic Lethality
FKH2 YNL068C Forkhead family transcription factor regulation 1 1 1 ChEC-Seq experiments
BUR6 YER159C Subunit of a heterodimeric NC2 transcription regulator complex regulation 1 1 1 chromatin immunoprecipitation-chip evidence
REB1 YBR049C RNA polymerase I enhancer binding protein regulation 1 1 1 chromatin immunoprecipitation-chip evidence
CDC73 YLR418C Component of the Paf1p complex genetic 1 1 1 Negative Genetic
DED81 YHR019C Cytosolic asparaginyl-tRNA synthetase genetic 1 1 1 Positive Genetic
CKB1 YGL019W Beta regulatory subunit of casein kinase 2 (CK2) genetic 1 1 1 Positive Genetic
RLP7 YNL002C Nucleolar protein similar to large ribosomal subunit L7 proteins physical 1 1 1 Affinity Capture-MS
RIC1 YLR039C Protein involved in retrograde transport to the cis-Golgi network genetic 1 1 1 Positive Genetic
PRP19 YLL036C Splicing factor associated with the spliceosome genetic 1 1 1 Negative Genetic
BRE1 YDL074C E3 ubiquitin ligase genetic 1 1 1 Synthetic Growth Defect
LOC1 YFR001W Nucleolar protein involved in the asymmetric localization of ASH1 mRNA genetic 1 1 1 Negative Genetic
RRP5 YMR229C RNA binding protein involved in synthesis of 18S and 5.8S rRNAs genetic 1 1 1 Negative Genetic
SAP185 YJL098W Protein that forms a complex with the Sit4p protein phosphatase physical 1 1 1 Affinity Capture-MS
PAT1 YCR077C Deadenylation-dependent mRNA-decapping factor genetic 1 1 1 Positive Genetic
LRP1 YHR081W Nuclear exosome-associated nucleic acid binding protein genetic 1 1 1 Negative Genetic
RPA49 YNL248C RNA polymerase I subunit A49 genetic 1 1 1 Negative Genetic
CBC2 YPL178W Small subunit of the heterodimeric cap binding complex with Sto1p genetic 1 1 1 Negative Genetic
THP2 YHR167W Subunit of the THO and TREX complexes genetic 1 1 1 Positive Genetic
GCD11 YER025W Gamma subunit of the translation initiation factor eIF2 genetic 1 1 1 Positive Genetic
SPT16 YGL207W Subunit of the heterodimeric FACT complex (Spt16p-Pob3p) physical 1 1 1 Affinity Capture-MS
CCR4 YAL021C Component of the CCR4-NOT transcriptional complex physical 1 1 1 Affinity Capture-RNA
LHP1 YDL051W RNA binding protein required for maturation of tRNA and U6 snRNA genetic 1 1 1 Negative Genetic
SUP35 YDR172W Translation termination factor eRF3 genetic 1 1 1 Positive Genetic
ADE4 YMR300C Phosphoribosylpyrophosphate amidotransferase (PRPPAT) genetic 1 1 1 Positive Genetic
CDC36 YDL165W Component of the CCR4-NOT core complex, involved in mRNA decapping genetic 1 1 1 Negative Genetic
DBP10 YDL031W Putative ATP-dependent RNA helicase of the DEAD-box protein family genetic 1 1 1 Negative Genetic
BUD31 YCR063W Component of the SF3b subcomplex of the U2 snRNP genetic 1 1 1 Positive Genetic
DCP2 YNL118C Catalytic subunit of Dcp1p-Dcp2p decapping enzyme complex genetic 1 1 1 Negative Genetic
FKS1 YLR342W Catalytic subunit of 1,3-beta-D-glucan synthase genetic 1 1 1 Positive Genetic
GCN1 YGL195W Positive regulator of the Gcn2p kinase activity genetic 1 1 1 Negative Genetic
DHH1 YDL160C Cytoplasmic DEAD-box helicase and mRNA decapping activator physical 1 1 1 Affinity Capture-RNA
CLU1 YMR012W Subunit of the eukaryotic translation initiation factor 3 (eIF3) genetic 1 1 1 Positive Genetic
KRS1 YDR037W Lysyl-tRNA synthetase genetic 1 1 1 Positive Genetic
HSC82 YMR186W Cytoplasmic chaperone of the Hsp90 family physical 1 1 1 Affinity Capture-MS
HRR25 YPL204W Conserved casein kinase, monopolin subunit physical 1 1 1 Biochemical Activity
MSH2 YOL090W Protein that binds to DNA mismatches genetic 1 1 1 Negative Genetic
MUM2 YBR057C Protein essential for meiotic DNA replication and sporulation genetic 1 1 1 Negative Genetic
NHP6B YBR089C-A High-mobility group (HMG) protein physical 1 1 1 PCA
ATG18 YFR021W Phosphoinositide binding protein genetic 1 1 1 Negative Genetic
NUP116 YMR047C FG-nucleoporin component of central core of the nuclear pore complex genetic 1 1 1 Negative Genetic
APM1 YPL259C Mu1-like medium subunit of the AP-1 complex physical 1 1 1 Affinity Capture-MS
DUR12 YBR208C Urea amidolyase genetic 1 1 1 Negative Genetic
POP7 YBR167C Subunit of RNase MRP, nuclear RNase P and telomerase genetic 1 1 1 Positive Genetic
PPH3 YDR075W Catalytic subunit of protein phosphatase PP4 complex genetic 1 1 1 Negative Genetic
PRE9 YGR135W Alpha 3 subunit of the 20S proteasome genetic 1 1 1 Positive Genetic
ILV1 YER086W Threonine deaminase, catalyzes first step in isoleucine biosynthesis genetic 1 1 1 Positive Genetic
RFA1 YAR007C Subunit of heterotrimeric Replication Protein A (RPA) physical 1 1 1 Two-hybrid
RPL2A YFR031C-A Ribosomal 60S subunit protein L2A genetic 1 1 1 Negative Genetic
SCS2 YER120W Integral ER membrane protein, regulates phospholipid metabolism genetic 1 1 1 Negative Genetic
SIN3 YOL004W Component of both the Rpd3S and Rpd3L histone deacetylase complexes genetic 1 1 1 Negative Genetic
RXT2 YBR095C Component of the histone deacetylase Rpd3L complex genetic 1 1 1 Negative Genetic
PRP9 YDL030W Subunit of the SF3a splicing factor complex genetic 1 1 1 Negative Genetic
SAK1 YER129W Upstream serine/threonine kinase for the SNF1 complex genetic 1 1 1 Negative Genetic
SNF1 YDR477W AMP-activated S/T protein kinase genetic 1 1 1 Negative Genetic
SNF5 YBR289W Subunit of the SWI/SNF chromatin remodeling complex genetic 1 1 1 Negative Genetic
ROM2 YLR371W Guanine nucleotide exchange factor (GEF) for Rho1p and Rho2p genetic 1 1 1 Negative Genetic
SEN54 YPL083C Subunit of the tRNA splicing endonuclease physical 1 1 1 Affinity Capture-MS
SBP1 YHL034C Protein that binds eIF4G and has a role in repression of translation physical 1 1 1 Affinity Capture-RNA
GTO3 YMR251W Omega class glutathione transferase genetic 1 1 1 Negative Genetic
SKN1 YGR143W Protein involved in sphingolipid biosynthesis genetic 1 1 1 Negative Genetic
DRS2 YAL026C Trans-golgi network aminophospholipid translocase (flippase) genetic 1 1 1 Negative Genetic
SKI3 YPR189W Ski complex component and TPR protein genetic 1 1 1 Negative Genetic
NGL2 YMR285C Protein involved in 5.8S rRNA processing genetic 1 1 1 Negative Genetic
MMS2 YGL087C Ubiquitin-conjugating enzyme variant genetic 1 1 1 Negative Genetic
RIM15 YFL033C Protein kinase involved in cell proliferation in response to nutrients genetic 1 1 1 Negative Genetic
ASA1 YPR085C Subunit of the ASTRA complex, involved in chromatin remodeling genetic 1 1 1 Positive Genetic
PBS2 YJL128C MAP kinase kinase of the HOG signaling pathway genetic 1 1 1 Negative Genetic
LCP5 YER127W Essential protein involved in maturation of 18S rRNA genetic 1 1 1 Positive Genetic
CAF120 YNL278W Part of the CCR4-NOT transcriptional regulatory complex genetic 1 1 1 Negative Genetic
CCT5 YJR064W Subunit of the cytosolic chaperonin Cct ring complex physical 1 1 1 Affinity Capture-MS
SLX5 YDL013W Subunit of the Slx5-Slx8 SUMO-targeted Ub ligase (STUbL) complex genetic 1 1 1 Positive Genetic
GPD2 YOL059W NAD-dependent glycerol 3-phosphate dehydrogenase genetic 1 1 1 Negative Genetic
SPH1 YLR313C Protein involved in shmoo formation and bipolar bud site selection genetic 1 1 1 Negative Genetic
ELM1 YKL048C Serine/threonine protein kinase genetic 1 1 1 Positive Genetic
NUP49 YGL172W FG-nucleoporin component of central core of the nuclear pore complex genetic 1 1 1 Negative Genetic
SEN15 YMR059W Subunit of the tRNA splicing endonuclease genetic 1 1 1 Negative Genetic
HOG1 YLR113W Mitogen-activated protein kinase involved in osmoregulation genetic 1 1 1 Negative Genetic
ISU1 YPL135W Conserved protein of the mitochondrial matrix genetic 1 1 1 Negative Genetic
HHO1 YPL127C Histone H1, linker histone with roles in meiosis and sporulation genetic 1 1 1 Negative Genetic
RAS2 YNL098C GTP-binding protein genetic 1 1 1 Positive Genetic
TDA1 YMR291W Protein kinase of unknown cellular role genetic 1 1 1 Negative Genetic
AIR1 YIL079C Zinc knuckle protein genetic 1 1 1 Negative Genetic
TPK3 YKL166C cAMP-dependent protein kinase catalytic subunit physical 1 1 1 Biochemical Activity
GID10 YGR066C Recognition component (N-recognin) of the Pro/N-degron pathway genetic 1 1 1 Negative Genetic
PTC6 YCR079W Mitochondrial type 2C protein phosphatase (PP2C) physical 1 1 1 Affinity Capture-MS
DAP1 YPL170W Heme-binding protein genetic 1 1 1 Negative Genetic
FMP48 YGR052W Putative protein of unknown function genetic 1 1 1 Negative Genetic
MCM16 YPR046W Component of the Ctf19 complex and the COMA subcomplex genetic 1 1 1 Positive Genetic
MSB2 YGR014W Mucin family member involved in various signaling pathways genetic 1 1 1 Negative Genetic
NDL1 YLR254C Homolog of nuclear distribution factor NudE genetic 1 1 1 Negative Genetic
ART5 YGR068C Protein proposed to regulate endocytosis of plasma membrane proteins genetic 1 1 1 Negative Genetic
GIP3 YPL137C Cytoplasmic protein that regulates protein phosphatase 1 Glc7p genetic 1 1 1 Negative Genetic
CHL1 YPL008W Probable DNA helicase genetic 1 1 1 Negative Genetic
NHP6A YPR052C High-mobility group (HMG) protein physical 1 1 1 PCA
SWR1 YDR334W Swi2/Snf2-related ATPase genetic 1 1 1 Positive Genetic
BDP1 YNL039W Essential subunit of RNA polymerase III transcription factor (TFIIIB) genetic 1 1 1 Negative Genetic
CBF1 YJR060W Basic helix-loop-helix (bHLH) protein genetic 1 1 1 Negative Genetic
RPS11B YBR048W Protein component of the small (40S) ribosomal subunit genetic 1 1 1 Negative Genetic
SSC1 YJR045C Hsp70 family ATPase physical 1 1 1 Affinity Capture-MS
BFR1 YOR198C Component of mRNP complexes associated with polyribosomes physical 1 1 1 Affinity Capture-RNA
MOT2 YER068W Ubiquitin-protein ligase subunit of the CCR4-NOT complex genetic 1 1 1 Negative Genetic
DBP5 YOR046C Cytoplasmic ATP-dependent RNA helicase of the DEAD-box family genetic 1 1 1 Synthetic Growth Defect
RPA14 YDR156W RNA polymerase I subunit A14 genetic 1 1 1 Positive Genetic
SEC61 YLR378C Conserved ER protein translocation channel physical 1 1 1 Affinity Capture-MS
SIP5 YMR140W Protein of unknown function genetic 1 1 1 Negative Genetic
SYN8 YAL014C Endosomal SNARE related to mammalian syntaxin 8 genetic 1 1 1 Negative Genetic
TEL1 YBL088C Protein kinase primarily involved in telomere length regulation genetic 1 1 1 Negative Genetic
TIM23 YNR017W Essential component of the TIM23 complex physical 1 1 1 Affinity Capture-MS
SRN2 YLR119W Component of the ESCRT-I complex genetic 1 1 1 Negative Genetic
EMP47 YFL048C Integral membrane component of ER-derived COPII-coated vesicles genetic 1 1 1 Negative Genetic
NUP170 YBL079W Subunit of inner ring of nuclear pore complex (NPC) genetic 1 1 1 Negative Genetic
SPT10 YJL127C Histone H3 acetylase with a role in transcriptional regulation physical 1 1 1 Affinity Capture-MS
EMC5 YIL027C Member of conserved ER transmembrane complex genetic 1 1 1 Negative Genetic
CAT8 YMR280C Zinc cluster transcriptional activator genetic 1 1 1 Negative Genetic
BPT1 YLL015W ABC type transmembrane transporter of MRP/CFTR family genetic 1 1 1 Positive Genetic
SEN2 YLR105C Subunit of the tRNA splicing endonuclease genetic 1 1 1 Negative Genetic
LRS4 YDR439W Nucleolar protein that forms a complex with Csm1p genetic 1 1 1 Positive Genetic
ALG5 YPL227C UDP-glucose:dolichyl-phosphate glucosyltransferase genetic 1 1 1 Negative Genetic
SUR1 YPL057C Mannosylinositol phosphorylceramide (MIPC) synthase catalytic subunit genetic 1 1 1 Negative Genetic
PHB1 YGR132C Subunit of the prohibitin complex (Phb1p-Phb2p) genetic 1 1 1 Negative Genetic
IXR1 YKL032C Transcriptional repressor that regulates hypoxic genes during normoxia genetic 1 1 1 Negative Genetic
EUC1 YMR111C Sequence-specific DNA binding protein genetic 1 1 1 Positive Genetic
ROG3 YFR022W Alpha-arrestin involved in ubiquitin-dependent endocytosis genetic 1 1 1 Negative Genetic
COQ9 YLR201C Protein required for ubiquinone biosynthesis and respiratory growth physical 1 1 1 Affinity Capture-MS
CSF1 YLR087C Protein with structural similarity to lipid transport protein Vps13p genetic 1 1 1 Positive Genetic
SRS2 YJL092W DNA helicase and DNA-dependent ATPase genetic 1 1 1 Dosage Lethality
TIM21 YGR033C Nonessential component of the TIM23 complex genetic 1 1 1 Negative Genetic
AGP2 YBR132C Plasma membrane regulator of polyamine and carnitine transport genetic 1 1 1 Negative Genetic
BAP3 YDR046C Amino acid permease genetic 1 1 1 Positive Genetic
MRH1 YDR033W Protein that localizes primarily to the plasma membrane genetic 1 1 1 Negative Genetic
NGR1 YBR212W RNA binding protein that negatively regulates growth rate genetic 1 1 1 Positive Genetic
PUF4 YGL014W Member of the PUF protein family physical 1 1 1 Affinity Capture-RNA
AEP3 YPL005W Peripheral mitochondrial inner membrane protein genetic 1 1 1 Positive Genetic
ADE16 YLR028C Enzyme of 'de novo' purine biosynthesis genetic 1 1 1 Negative Genetic
IZH3 YLR023C Membrane protein involved in zinc ion homeostasis genetic 1 1 1 Negative Genetic
ADE12 YNL220W Adenylosuccinate synthase physical 1 1 1 Affinity Capture-MS
PFK26 YIL107C 6-phosphofructo-2-kinase genetic 1 1 1 Negative Genetic
RPS0B YLR048W Protein component of the small (40S) ribosomal subunit genetic 1 1 1 Negative Genetic
ARV1 YLR242C Cortical ER protein genetic 1 1 1 Negative Genetic
PSP1 YDR505C Asn and gln rich protein of unknown function genetic 1 1 1 Negative Genetic
COQ8 YGL119W ATPase required for ubiquinone biosynthesis and respiratory growth physical 1 1 1 Affinity Capture-MS
PUF2 YPR042C PUF family mRNA-binding protein physical 1 1 1 Affinity Capture-RNA
BNA2 YJR078W Tryptophan 2,3-dioxygenase or indoleamine 2,3-dioxygenase genetic 1 1 1 Negative Genetic
SUB1 YMR039C Transcriptional regulator genetic 1 1 1 Negative Genetic
DOT5 YIL010W Nuclear thiol peroxidase genetic 1 1 1 Negative Genetic
PSR2 YLR019W Putative plasma membrane phosphatase genetic 1 1 1 Negative Genetic
EEB1 YPL095C Acyl-coenzymeA:ethanol O-acyltransferase genetic 1 1 1 Negative Genetic
EAR1 YMR171C Specificity factor required for Rsp5p-dependent ubiquitination genetic 1 1 1 Negative Genetic
LGE1 YPL055C Protein involved in histone H2B ubiquitination genetic 1 1 1 Synthetic Growth Defect
COS12 YGL263W Endosomal protein involved in turnover of plasma membrane proteins genetic 1 1 1 Negative Genetic
TPO2 YGR138C Polyamine transporter of the major facilitator superfamily genetic 1 1 1 Negative Genetic
AVL9 YLR114C Conserved protein involved in exocytic transport from the Golgi genetic 1 1 1 Negative Genetic
COX20 YDR231C Mitochondrial inner membrane protein physical 1 1 1 Affinity Capture-MS
RCE1 YMR274C Type II CAAX prenyl protease genetic 1 1 1 Negative Genetic
IFM1 YOL023W Mitochondrial translation initiation factor 2 genetic 1 1 1 Positive Genetic
EMC10 YDR056C Putative protein of unknown function genetic 1 1 1 Negative Genetic
MIC10 YCL057C-A Conserved component of the MICOS complex physical 1 1 1 Affinity Capture-MS
NPR3 YHL023C Subunit of the Iml1p/SEACIT complex genetic 1 1 1 Negative Genetic
AIM36 YMR157C Protein of unknown function genetic 1 1 1 Negative Genetic
MGR2 YPL098C Subunit of the TIM23 translocase complex genetic 1 1 1 Negative Genetic
SEI1 YLR404W Seipin involved in lipid droplet (LD) assembly genetic 1 1 1 Negative Genetic
HMI1 YOL095C Mitochondrial inner membrane localized ATP-dependent DNA helicase genetic 1 1 1 Negative Genetic
MME1 YMR166C Transporter of the mitochondrial inner membrane that exports magnesium genetic 1 1 1 Positive Genetic
QRI7 YDL104C Protein involved in threonylcarbamoyl adenosine biosynthesis genetic 1 1 1 Negative Genetic
OXR1 YPL196W Protein involved in regulation of vacuolar V-ATPase genetic 1 1 1 Negative Genetic
PDH1 YPR002W Putative 2-methylcitrate dehydratase genetic 1 1 1 Negative Genetic
IRS4 YKR019C EH domain-containing protein genetic 1 1 1 Negative Genetic
SPS2 YDR522C Protein expressed during sporulation genetic 1 1 1 Negative Genetic
RNP1 YLL046C Ribonucleoprotein that contains two RNA recognition motifs (RRM) genetic 1 1 1 Negative Genetic
UTP4 YDR324C Subunit of U3-containing 90S preribosome and SSU processome complexes genetic 1 1 1 Negative Genetic
YPT6 YLR262C Rab family GTPase genetic 1 1 1 Positive Genetic
VMA21 YGR105W Integral membrane protein required for V-ATPase function genetic 1 1 1 Synthetic Growth Defect
YRB30 YGL164C RanGTP-binding protein genetic 1 1 1 Positive Genetic
YDR061W YDR061W Protein with similarity to ABC transporter family members genetic 1 1 1 Negative Genetic
UBC13 YDR092W E2 ubiquitin-conjugating enzyme genetic 1 1 1 Negative Genetic
YGR017W YGR017W Putative protein of unknown function genetic 1 1 1 Negative Genetic
YGR018C YGR018C Protein of unknown function genetic 1 1 1 Negative Genetic
VPS53 YJL029C Component of the GARP (Golgi-associated retrograde protein) complex genetic 1 1 1 Negative Genetic
YLR225C YLR225C Putative protein of unknown function genetic 1 1 1 Negative Genetic
VPS38 YLR360W Part of a Vps34p phosphatidylinositol 3-kinase complex genetic 1 1 1 Negative Genetic
YMR253C YMR253C Putative protein of unknown function genetic 1 1 1 Negative Genetic
YNL046W YNL046W Putative protein of unknown function genetic 1 1 1 Negative Genetic
ULA1 YPL003W Protein that activates Rub1p (NEDD8) before neddylation genetic 1 1 1 Negative Genetic
YDC1 YPL087W Alkaline dihydroceramidase, involved in sphingolipid metabolism genetic 1 1 1 Negative Genetic
VPS30 YPL120W Subunit of phosphatidylinositol (PtdIns) 3-kinase complexes I and II genetic 1 1 1 Negative Genetic
UME1 YPL139C Component of both the Rpd3S and Rpd3L histone deacetylase complexes genetic 1 1 1 Negative Genetic
YPL247C YPL247C Putative protein of unknown function genetic 1 1 1 Negative Genetic
YPR127W YPR127W Putative pyridoxine 4-dehydrogenase genetic 1 1 1 Negative Genetic
VPS4 YPR173C AAA-ATPase involved in multivesicular body (MVB) protein sorting genetic 1 1 1 Negative Genetic
SUC2 YIL162W Invertase genetic 1 1 1 Negative Genetic
TRM44 YPL030W tRNA(Ser) Um(44) 2'-O-methyltransferase genetic 1 1 1 Negative Genetic