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Ubiquitin-60S ribosomal protein L40



Drosophila melanogaster (Fruit fly)
Reviewed-Annotation score: Annotation score: 5 out of 5-Experimental evidence at protein leveli


Ubiquitin: exists either covalently attached to another protein, or free (unanchored). When covalently bound, it is conjugated to target proteins via an isopeptide bond either as a monomer (monoubiquitin), a polymer linked via different Lys residues of the ubiquitin (polyubiquitin chains) or a linear polymer linked via the initiator Met of the ubiquitin (linear polyubiquitin chains). Polyubiquitin chains, when attached to a target protein, have different functions depending on the Lys residue of the ubiquitin that is linked: Lys-48-linked is involved in protein degradation via the proteasome. Linear polymer chains formed via attachment by the initiator Met lead to cell signaling. Ubiquitin is usually conjugated to Lys residues of target proteins, however, in rare cases, conjugation to Cys or Ser residues has been observed. When polyubiquitin is free (unanchored-polyubiquitin), it also has distinct roles, such as in activation of protein kinases, and in signaling (By similarity).By similarity
60S ribosomal protein L40: component of the 60S subunit of the ribosome.By similarity


In Drosophila ubiquitin is encoded by 3 different genes. RpL40 and RpS27A genes code for a single copy of ubiquitin fused to the ribosomal proteins L40 and S27a, respectively. Ubi-p63E gene codes for a polyubiquitin precursor with 10 exact head to tail repeats.
For a better understanding, features related to ubiquitin are only indicated for the first chain.


Feature keyPosition(s)DescriptionActionsGraphical viewLength
Binding sitei54Activating enzyme1
Sitei68Essential for function1
Binding sitei72Activating enzyme1

GO - Molecular functioni

  • protein tag Source: FlyBase
  • structural constituent of ribosome Source: FlyBase

GO - Biological processi

  • cellular protein modification process Source: FlyBase
  • cytoplasmic translation Source: FlyBase
  • Notch signaling pathway Source: Reactome
  • protein ubiquitination Source: FlyBase
  • translation Source: FlyBase
  • ubiquitin-dependent protein catabolic process Source: FlyBase


Molecular functionRibonucleoprotein, Ribosomal protein

Enzyme and pathway databases

ReactomeiR-DME-110312. Translesion synthesis by REV1.
R-DME-110320. Translesion Synthesis by POLH.
R-DME-1169091. Activation of NF-kappaB in B cells.
R-DME-1234176. Oxygen-dependent proline hydroxylation of Hypoxia-inducible Factor Alpha.
R-DME-1253288. Downregulation of ERBB4 signaling.
R-DME-1295596. Spry regulation of FGF signaling.
R-DME-1358803. Downregulation of ERBB2:ERBB3 signaling.
R-DME-156827. L13a-mediated translational silencing of Ceruloplasmin expression.
R-DME-174048. APC/C:Cdc20 mediated degradation of Cyclin B.
R-DME-174084. Autodegradation of Cdh1 by Cdh1:APC/C.
R-DME-174113. SCF-beta-TrCP mediated degradation of Emi1.
R-DME-174178. APC/C:Cdh1 mediated degradation of Cdc20 and other APC/C:Cdh1 targeted proteins in late mitosis/early G1.
R-DME-174184. Cdc20:Phospho-APC/C mediated degradation of Cyclin A.
R-DME-179409. APC-Cdc20 mediated degradation of Nek2A.
R-DME-1799339. SRP-dependent cotranslational protein targeting to membrane.
R-DME-182971. EGFR downregulation.
R-DME-187577. SCF(Skp2)-mediated degradation of p27/p21.
R-DME-195253. Degradation of beta-catenin by the destruction complex.
R-DME-201681. TCF dependent signaling in response to WNT.
R-DME-209360. Ubiquitination and proteolysis of phosphorylated CI.
R-DME-209447. Activation of the IkappaB kinase complex, KEY:IRD5 dimer:KEY.
R-DME-209461. Ubiquitination and degradation of phosphorylated ARM.
R-DME-209560. NF-kB is activated and signals survival.
R-DME-2122948. Activated NOTCH1 Transmits Signal to the Nucleus.
R-DME-2173788. Downregulation of TGF-beta receptor signaling.
R-DME-2173791. TGF-beta receptor signaling in EMT (epithelial to mesenchymal transition).
R-DME-2173795. Downregulation of SMAD2/3:SMAD4 transcriptional activity.
R-DME-2173796. SMAD2/SMAD3:SMAD4 heterotrimer regulates transcription.
R-DME-2559582. Senescence-Associated Secretory Phenotype (SASP).
R-DME-3769402. Deactivation of the beta-catenin transactivating complex.
R-DME-432395. Degradation of TIM.
R-DME-432524. Degradation of PER.
R-DME-446652. Interleukin-1 family signaling.
R-DME-450302. activated TAK1 mediates p38 MAPK activation.
R-DME-450321. JNK (c-Jun kinases) phosphorylation and activation mediated by activated human TAK1.
R-DME-450408. AUF1 (hnRNP D0) binds and destabilizes mRNA.
R-DME-4608870. Asymmetric localization of PCP proteins.
R-DME-4641257. Degradation of AXIN.
R-DME-4641258. Degradation of DVL.
R-DME-5205685. Pink/Parkin Mediated Mitophagy.
R-DME-532668. N-glycan trimming in the ER and Calnexin/Calreticulin cycle.
R-DME-5358346. Hedgehog ligand biogenesis.
R-DME-538864. Degradation of CRY.
R-DME-5607761. Dectin-1 mediated noncanonical NF-kB signaling.
R-DME-5610785. GLI3 is processed to GLI3R by the proteasome.
R-DME-5632684. Hedgehog 'on' state.
R-DME-5654726. Negative regulation of FGFR1 signaling.
R-DME-5654727. Negative regulation of FGFR2 signaling.
R-DME-5654732. Negative regulation of FGFR3 signaling.
R-DME-5654733. Negative regulation of FGFR4 signaling.
R-DME-5655862. Translesion synthesis by POLK.
R-DME-5656121. Translesion synthesis by POLI.
R-DME-5656169. Termination of translesion DNA synthesis.
R-DME-5675221. Negative regulation of MAPK pathway.
R-DME-5675482. Regulation of necroptotic cell death.
R-DME-5676590. NIK-->noncanonical NF-kB signaling.
R-DME-5689603. UCH proteinases.
R-DME-5689877. Josephin domain DUBs.
R-DME-5689880. Ub-specific processing proteases.
R-DME-5689896. Ovarian tumor domain proteases.
R-DME-5689901. Metalloprotease DUBs.
R-DME-5693565. Recruitment and ATM-mediated phosphorylation of repair and signaling proteins at DNA double strand breaks.
R-DME-5696394. DNA Damage Recognition in GG-NER.
R-DME-5696395. Formation of Incision Complex in GG-NER.
R-DME-6781823. Formation of TC-NER Pre-Incision Complex.
R-DME-6782135. Dual incision in TC-NER.
R-DME-6782210. Gap-filling DNA repair synthesis and ligation in TC-NER.
R-DME-6807004. Negative regulation of MET activity.
R-DME-68949. Orc1 removal from chromatin.
R-DME-69017. CDK-mediated phosphorylation and removal of Cdc6.
R-DME-69229. Ubiquitin-dependent degradation of Cyclin D1.
R-DME-69231. Cyclin D associated events in G1.
R-DME-69601. Ubiquitin Mediated Degradation of Phosphorylated Cdc25A.
R-DME-72689. Formation of a pool of free 40S subunits.
R-DME-72706. GTP hydrolysis and joining of the 60S ribosomal subunit.
R-DME-8849469. PTK6 Regulates RTKs and Their Effectors AKT1 and DOK1.
R-DME-8854050. FBXL7 down-regulates AURKA during mitotic entry and in early mitosis.
R-DME-8856825. Cargo recognition for clathrin-mediated endocytosis.
R-DME-8856828. Clathrin-mediated endocytosis.
R-DME-8863795. Downregulation of ERBB2 signaling.
R-DME-8866652. Synthesis of active ubiquitin: roles of E1 and E2 enzymes.
R-DME-8866654. E3 ubiquitin ligases ubiquitinate target proteins.
R-DME-8941858. Regulation of RUNX3 expression and activity.
R-DME-8948747. Regulation of PTEN localization.
R-DME-8948751. Regulation of PTEN stability and activity.
R-DME-901032. ER Quality Control Compartment (ERQC).
R-DME-9010553. Regulation of expression of SLITs and ROBOs.
R-DME-912631. Regulation of signaling by CBL.
R-DME-917729. Endosomal Sorting Complex Required For Transport (ESCRT).
R-DME-936440. Negative regulators of DDX58/IFIH1 signaling.
R-DME-937039. IRAK1 recruits IKK complex.
R-DME-937042. IRAK2 mediated activation of TAK1 complex.
R-DME-975110. TRAF6 mediated IRF7 activation in TLR7/8 or 9 signaling.
R-DME-975144. IRAK1 recruits IKK complex upon TLR7/8 or 9 stimulation.
R-DME-975163. IRAK2 mediated activation of TAK1 complex upon TLR7/8 or 9 stimulation.
R-DME-975956. Nonsense Mediated Decay (NMD) independent of the Exon Junction Complex (EJC).
R-DME-975957. Nonsense Mediated Decay (NMD) enhanced by the Exon Junction Complex (EJC).
R-DME-983168. Antigen processing: Ubiquitination & Proteasome degradation.

Names & Taxonomyi

Protein namesi
Recommended name:
Ubiquitin-60S ribosomal protein L40
Alternative name(s):
Cleaved into the following 2 chains:
Gene namesi
Synonyms:Ubi-f, UBI-F52
ORF Names:CG2960
OrganismiDrosophila melanogaster (Fruit fly)
Taxonomic identifieri7227 [NCBI]
Taxonomic lineageiEukaryotaMetazoaEcdysozoaArthropodaHexapodaInsectaPterygotaNeopteraHolometabolaDipteraBrachyceraMuscomorphaEphydroideaDrosophilidaeDrosophilaSophophora
  • UP000000803 Componenti: Chromosome 2L

Organism-specific databases

FlyBaseiFBgn0003941. RpL40.

Subcellular locationi

Extracellular region or secreted Cytosol Plasma membrane Cytoskeleton Lysosome Endosome Peroxisome ER Golgi apparatus Nucleus Mitochondrion Manual annotation Automatic computational assertionGraphics by Christian Stolte; Source: COMPARTMENTS

Keywords - Cellular componenti

Cytoplasm, Nucleus

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
ChainiPRO_00003964421 – 76UbiquitinAdd BLAST76
ChainiPRO_000039644377 – 12860S ribosomal protein L40Add BLAST52

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Cross-linki48Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in ubiquitin)
Cross-linki76Glycyl lysine isopeptide (Gly-Lys) (interchain with K-? in acceptor proteins)

Keywords - PTMi

Isopeptide bond, Ubl conjugation

Proteomic databases



Gene expression databases

ExpressionAtlasiP18101. differential.
GenevisibleiP18101. DM.


Subunit structurei

60S ribosomal protein is part of the 60S ribosomal subunit.By similarity

Protein-protein interaction databases

BioGridi59826. 18 interactors.
IntActiP18101. 2 interactors.


3D structure databases

Select the link destinations:
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
4V6Welectron microscopy6.00Cm77-128[»]

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Domaini1 – 76Ubiquitin-likePROSITE-ProRule annotationAdd BLAST76

Sequence similaritiesi

In the N-terminal section; belongs to the ubiquitin family.Curated
In the C-terminal section; belongs to the eukaryotic ribosomal protein eL40 family.Curated

Phylogenomic databases

eggNOGiKOG0003. Eukaryota.
COG1552. LUCA.
COG5272. LUCA.

Family and domain databases

InterProiView protein in InterPro
IPR001975. Ribosomal_L40e.
IPR019956. Ubiquitin.
IPR029071. Ubiquitin-like_domsf.
IPR019954. Ubiquitin_CS.
IPR000626. Ubiquitin_dom.
PfamiView protein in Pfam
PF01020. Ribosomal_L40e. 1 hit.
PF00240. ubiquitin. 1 hit.
SMARTiView protein in SMART
SM01377. Ribosomal_L40e. 1 hit.
SM00213. UBQ. 1 hit.
SUPFAMiSSF54236. SSF54236. 1 hit.
PROSITEiView protein in PROSITE
PS00299. UBIQUITIN_1. 1 hit.
PS50053. UBIQUITIN_2. 1 hit.


Sequence statusi: Complete.

Sequence processingi: The displayed sequence is further processed into a mature form.

P18101-1 [UniParc]FASTAAdd to basket

« Hide

        10         20         30         40         50
60 70 80 90 100
110 120
Mass (Da):14,729
Last modified:August 10, 2010 - v2

Sequence databases

Select the link destinations:
Links Updated
X53059 mRNA. Translation: CAA37227.1.
X59943 Genomic DNA. Translation: CAA42568.1.
AE014134 Genomic DNA. Translation: AAF51034.1.
RefSeqiNP_001260018.1. NM_001273089.2.
NP_476776.1. NM_057428.4.

Genome annotation databases

EnsemblMetazoaiFBtr0077470; FBpp0077159; FBgn0003941.
FBtr0334787; FBpp0306837; FBgn0003941.

Similar proteinsi

Entry informationi

Entry nameiRL40_DROME
AccessioniPrimary (citable) accession number: P18101
Secondary accession number(s): P68198
, Q0E8I1, Q9VKW6, Q9VQX7, Q9VZL4
Entry historyiIntegrated into UniProtKB/Swiss-Prot: November 1, 1990
Last sequence update: August 10, 2010
Last modified: November 22, 2017
This is version 134 of the entry and version 2 of the sequence. See complete history.
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programDrosophila annotation project


Keywords - Technical termi

3D-structure, Complete proteome, Reference proteome


  1. Drosophila
    Drosophila: entries, gene names and cross-references to FlyBase
  2. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
  3. Ribosomal proteins
    Ribosomal proteins families and list of entries
  4. SIMILARITY comments
    Index of protein domains and families