Stress granule

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Stress granule dynamics

In the cellular biology, stress granules are biomolecular condensates in the cytosol composed of proteins and RNA that assemble into 0.1–2 μm membraneless organelles when the cell is under stress.[1][2] The mRNA molecules found in stress granules are stalled translation pre-initiation complexes associated with 40S ribosomal subunits, translation initiation factors, poly(A)+ mRNA and RNA-binding proteins (RBPs). While they are membraneless organelles, stress granules have been proposed to be associated with the endoplasmatic reticulum.[3] There are also nuclear stress granules. This article is about the cytosolic variety.

Proposed functions

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The function of stress granules remains largely unknown. Stress granules have long been proposed to have a function to protect RNA from harmful conditions, thus their appearance under stress.[4] The accumulation of RNA into dense globules could keep them from reacting with harmful chemicals and safeguard the information coded in their RNA sequence.

Stress granules might also function as a decision point for untranslated mRNA. Molecules can go down one of three paths: further storage, degradation, or re-initiation of translation.[5] Conversely, it has also been argued that stress granules are not important sites for mRNA storage nor do they serve as an intermediate location for mRNA in transit between a state of storage and a state of degradation.[6]

Efforts to identify all RNA within stress granules (the stress granule transcriptome) in an unbiased way by sequencing RNA from biochemically purified stress granule "cores" have shown that RNA are not recruited to stress granules in a sequence-specific manner, but rather generically, with longer and/or less-optimally translated transcripts being enriched.[7] These data imply that the stress granule transcriptome is influenced by the valency of RNA (for proteins or other RNA) and by the rates of RNA run-off from polysomes. The latter is further supported by recent single molecule imaging studies.[8] Furthermore, it was estimated that only about 15% of the total mRNA in the cell is localized to stress granules,[7] suggesting that stress granules only influence a minority of mRNA in the cell and may not be as important for mRNA processing as previously thought.[7][9] That said, these studies represent only a snapshot in time, and it is likely that a larger fraction of mRNA are at one point stored in stress granules due to those RNA transiting in and out.

The stress proteins that are the main component of stress granules in plant cells are molecular chaperones that sequester, protect, and possibly repair proteins that unfold during heat and other types of stress.[10][11] Therefore, any association of mRNA with stress granules may simply be a side effect of the association of partially unfolded RNA-binding proteins with stress granules,[12] similar to the association of mRNA with proteasomes.[13]

Formation

[edit]

Environmental stressors trigger cellular signaling, eventually leading to the formation of stress granules. In vitro, these stressors can include heat, cold, oxidative stress (sodium arsenite), endoplasmic reticulum stress (thapsigargin), proteasome inhibition (MG132), hyperosmotic stress, ultraviolet radiation, inhibition of eIF4A (pateamine A, hippuristanol, or RocA), nitric oxide accumulation after treatment with 3-morpholinosydnonimine (SIN-1),[14] perturbation of pre-mRNA splicing,[15] and other stressors, like puromycin, which result in disassembled polysomes.[16] Many of these stressors result in the activation of particular stress-associated kinases (HRI, PERK, PKR, and GCN2), translational inhibition and stress granule formation.[16] Stress granules will also form upon Gαq activation in a mechanism that involves the release of stress granule associated proteins from the cytosolic population of the Gαq effector phospholipase Cβ.[17]

Stress granule formation is often downstream of the stress-activated phosphorylation of eukaryotic translation initiation factor eIF2α; this does not hold true for all types of stressors that induce stress granules,[16] for instance, eIF4A inhibition. Further downstream, prion-like aggregation of the protein TIA-1 promotes the formation of stress granules. The term prion-like is used because aggregation of TIA-1 is concentration dependent, inhibited by chaperones, and because the aggregates are resistant to proteases.[18] It has also been proposed that microtubules play a role in the formation of stress granules, perhaps by transporting granule components. This hypothesis is based on the fact that disruption of microtubules with the chemical nocodazole blocks the appearance of the granules.[19] Furthermore, many signaling molecules have been shown to regulate the formation or dynamics of stress granules; these include the "master energy sensor" AMP-activated protein kinase (AMPK),[20] the O-GlcNAc transferase enzyme (OGT),[21] and the pro-apoptotic kinase ROCK1.[22]

Potential roles of RNA-RNA interactions

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RNA phase transitions driven in part by intermolecular RNA-RNA interactions may play a role in stress granule formation. Similar to intrinsically disordered proteins, total RNA extracts are capable of undergoing phase separation in physiological conditions in vitro.[23] RNA-seq analyses demonstrate that these assemblies share a largely overlapping transcriptome with stress granules,[23][7] with RNA enrichment in both being predominately based on the length of the RNA. Further, stress granules contain many RNA helicases,[24] including the DEAD/H-box helicases Ded1p/DDX3, eIF4A1, and RHAU.[25] In yeast, catalytic ded1 mutant alleles give rise to constitutive stress granules[26] ATPase-deficient DDX3X (the mammalian homolog of Ded1) mutant alleles are found in pediatric medulloblastoma,[27] and these coincide with constitutive granular assemblies in patient cells.[28] These mutant DDX3 proteins promote stress granule assembly in HeLa cells.[28] In mammalian cells, RHAU mutants lead to reduced stress granule dynamics.[25] Thus, some hypothesize that RNA aggregation facilitated by intermolecular RNA-RNA interactions plays a role in stress granule formation, and that this role may be regulated by RNA helicases.[29] There is also evidence that RNA within stress granules is more compacted, compared to RNA in the cytoplasm, and that the RNA is found to be post-translationally modified by N6-methyladenosine (m6A) on its 5' ends or RNA acetylation ac4C.[30][31][32] Recent work has shown that the highly abundant translation initiation factor and DEAD-box protein eIF4A limits stress granule formation. It does so through its ability to bind ATP and RNA, acting analogously to protein chaperones like Hsp70.[33]

Connection with processing bodies

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Stress granules and P-bodies (processing bodies) share RNA and protein components, both appear under stress, and can physically associate with one another. As of 2018, of the ~660 proteins identified as localizing to stress granules, ~11% also have been identified as processing body-localized proteins (see below). The protein G3BP1 is necessary for the proper docking of processing bodies and stress granules to each other, which may be important for the preservation of polyadenylated mRNA.[34]

Although some protein components are shared between stress granules and processing bodies, the majority of proteins in either structure are uniquely localized to either structure.[35] While both stress granules and P-bodies are associated with mRNA, processing bodies have been long proposed to be sites of mRNA degradation because they contain enzymes such as DCP1/2 and XRN1 that are known to degrade mRNA.[36] However, others have demonstrated that mRNA associated with processing bodies are largely translationally repressed but not degraded.[35] It has also been proposed that mRNA selected for degradation are passed from stress granules to processing bodies,[36] though there is also data suggesting that processing bodies precede and promote stress granule formation.[37]

Protein composition of stress granules

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The complete proteome of stress granules is still unknown, but efforts have been made to catalog all of the proteins that have been experimentally demonstrated to transit into stress granules.[38][39][40] Importantly, different stressors can result in stress granules with different protein components.[16] Many stress granule-associated proteins have been identified by transiently stressing cultured cells and utilizing microscopy to detect the localization of a protein of interest either by expressing that protein fused to a fluorescent protein (i.e. green fluorescent protein (GFP)) and/or by fixing cells and using antibodies to detect the protein of interest along with known protein markers of stress granules (immunocytochemistry).[41]

In 2016, stress granule "cores" were experimentally identified and then biochemically purified for the first time. Proteins in the cores were identified in an unbiased manner using mass spectrometry. This technical advance lead to the identification of hundreds of new stress granule-localized proteins.[42][24][43]

The proteome of stress granules has also been experimentally determined by using two slightly different proximity labeling approaches. One of these proximity labeling approaches is the ascorbate peroxidase (APEX) method, in which cells are engineered to express a known stress granule protein, such as G3BP1, fused to a modified ascorbate peroxidase enzyme called APEX.[38][44] Upon incubating the cells in biotin and treating the cells with hydrogen peroxide, the APEX enzyme will be briefly activated to biotinylate all proteins in close proximity to the protein of interest, in this case G3BP1 within stress granules. Proteins that are biotinylated can then be isolated via streptavidin and identified using mass spectrometry. The APEX technique was used to identify ~260 stress granule-associated proteins in several cell types, including neurons, and with various stressors. Of the 260 proteins identified in this study, ~143 had not previously been demonstrated to be stress granule-associated.[44]

Another proximity labeling method used to determine the proteome of stress granules is BioID.[45] BioID is similar to the APEX approach, in that a biotinylating protein (BirA* instead of APEX) was expressed in cells as a fusion protein with several known stress granule-associated proteins. Proteins in close proximity to BirA* will be biotinylated and are then identified by mass spectrometry. Youn et al. used this method to identify/predict 138 proteins as stress granule-associated and 42 as processing body-associated.[45]

A curated database of stress granule-associated proteins can be found here [1].[40]

The following is a list of proteins that have been demonstrated to localize to stress granules (compiled from [38][39][24][44][45][46]):

Gene ID Protein Name Description References Also found in processing bodies?
ABCF1 ABCF1 ATP Binding Cassette Subfamily F Member 1 [44]
ABRACL ABRACL ABRA C-Terminal Like [44]
ACAP1 ACAP1 ArfGAP With Coiled-Coil, Ankyrin Repeat And PH Domains 1 [44]
ACBD5 ACBD5 Acyl-CoA Binding Domain Containing 5 [44]
ACTBL2 ACTBL2 Beta-actin-like protein 2 [24] yes[35]
ACTR1A ACTR1A Alpha-centractin [24]
ACTR1B ACTR1B Beta-centractin [24]
ADAR ADAR1 Adenosine Deaminase, RNA Specific [47][24]
ADD1 Adducin 1 Adducin 1 [44]
AGO1 Argonaute 1/EIF2C1 Argonaute 1, RISC Catalytic Component [44][48] yes[35]
AGO2 Argonaute 2 Argonaute 2, RISC Catalytic Component [44][49][48][50][24][51][46] yes[35]
AKAP8 AKAP8 A-Kinase Anchoring Protein 8 [46]
AKAP9 AKAP350 A-Kinase Anchoring Protein 9 [52]
AKAP13 AKAP13/LBC A-Kinase Anchoring Protein 13 [44][46]
ALDH18A1 ALDH18A1 Delta-1-pyrroline-5-carboxylate synthase [24]
ALG13 ALG13 ALG13, UDP-N-Acetylglucosaminyltransferase Subunit [45]
ALPK2 ALPK2/HAK Alpha Kinase 2 [46]
AMOTL2 AMOTL2/LCCP Angiomotin Like 2 [46]
ANKHD1 ANKHD1 Ankyrin Repeat and KH Domain Containing 1 [45] yes[45]
ANKRD17 ANKRD17/MASK2/GTAR Ankyrin Repeat Domain 17 [44][45] yes[45]
ANG Angiogenin Angiogenin [53]
ANP32E ANP32E Acidic leucine-rich nuclear phosphoprotein 32 family member E [24]
ANXA1 ANXA1 Annexin A1 [24]
ANXA11 ANXA11 Annexin 11 [44]
ANXA6 ANXA6 Annexin 6 [24]
ANXA7 ANXA7 Annexin 7 [24][44]
APEX1 APEX1 DNA-(apurinic or apyrimidinic site) lyase [24]
APOBEC3C APOBEC3C Apolipoprotein B mRNA Editing Enzyme Catalytic Subunit 3C [44][46]
APOBEC3G APOBEC3G Apolipoprotein B mRNA Editing Enzyme Catalytic Subunit 3G [48]
ARID2 ARID2/BAF200 AT-Rich Interaction Domain 2 [46]
ARPC1B ARPC1B Actin-related protein 2/3 complex subunit 1B [24]
AHSA1 AHA1 Activator Of HSP90 ATPase Activity 1 [54]
AQR AQR/IBP160 Aquarius Intron-Binding Spliceosomal Factor [44]
ARMC6 ARMC6 Armadillo Repeat Containing 6 [44]
ASCC1 ASCC1 Activating Signal Cointegrator 1 Complex Subunit 1 [44][45]
ASCC3 ASCC3 Activating Signal Cointegrator 1 Complex Subunit 3 [45]
ATAD2 ATAD2 ATPase family AAA domain-containing protein 2 [24]
ATAD3A ATAD3A ATPase family AAA domain-containing protein 3A [24] yes[35]
ATG3 ATG3 Autophagy Related 3 [44]
ATP5A1 ATP5A1 ATP synthase subunit alpha, mitochondrial [24]
ATP6V1G1 ATP6V1G1/ATP6G ATPase H+ Transporting V1 Subunit G1 [44]
ATXN2 Ataxin 2 Ataxin 2 [24][44][45][46][55][56][57][58][59][60]
ATXN2L Ataxin-2 like Ataxin 2 Like [24][44][45][46][57][60]
BAG3 BAG3 BAG family molecular chaperone regulator 3 [24]
BANF1 BANF1 Barrier-to-autointegration factor [24]
BAZ1B BAZ1B Bromodomain Adjacent To Zinc Finger Domain 1B [46]
BAZ2A BAZ2A Bromodomain Adjacent To Zinc Finger Domain 2A [46]
BCCIP BCCIP BRCA2 And CDKN1A Interacting Protein [44]
BCLAF1 BCLAF1 BCL2 Associated Transcription Factor 1 [44]
BICC1 BICC1 BicC Family RNA Binding Protein 1 [45]
BIRC2 BIRC2/CIAP1 Baculoviral IAP Repeat Containing 2 [46]
BLM BLM BLM RecQ Like Helicase [46]
BOD1L1 BOD1L1/FAM44A Biorientation Of Chromosomes In Cell Division 1 Like 1 [46]
BOLL BOULE Boule Homolog, RNA Binding Protein [61]
BRAT1 BRAT1 BRCA1-associated ATM activator 1 [24]
BRF1 BRF1 BRF1, RNA Polymerase III Transcription Initiation Factor Subunit [36]
BTG3 BTG3 BTG Anti-Proliferation Factor 3 [45] yes[45]
C9orf72 C9orf72 Uncharacterized protein C9orf72 [62][63]
C15orf52 C15orf52 Uncharacterized protein C15orf52 [24]
C20orf27 C20orf72 Chromosome 20 Open Reading Frame 27 [44]
C2CD3 C2CD3 C2 Calcium Dependent Domain Containing 3 [44]
CALML5 CALML5 Calmodulin-like protein 5 [24]
CALR Calreticulin/CRT Calreticulin [64]
CAMSAP1 CAMSAP1 Calmodulin Regulated Spectrin Associated Protein 1 [46]
CAP1 CAP1 Adenylyl cyclase-associated protein 1 [24]
CAPRIN1 Caprin-1 Cell Cycle Associated Protein 1 [44][45][65][52][66][24][67][34][68][60][46]
CAPZA2 CAPZA2 F-actin-capping protein subunit alpha-2 [24]
CAPZB CAPZB Capping Actin Protein Of Muscle Z-Line Subunit Beta [46]
CARHSP1 CARHSP1 Calcium-regulated heat stable protein 1 [24]
CASC3 MLN51/BTZ Cancer Susceptibility 3 [44][45][46][69][70]
CBFB CBFB Core-binding factor subunit beta [24]
CBS CBS Cystathionine Beta-Synthase [46]
CBX1 CBX1 Chromobox protein homolog 1 [24][60]
CBX3 CBX3 Chromobox protein homolog 3 [46]
CCAR1 CARP-1 Cell Division Cycle and Apoptosis Regulator 1 [52][46]
CCDC9 CCDC9 Coiled-Coil Domain Containing 9 [46]
CCDC9B CCDC9B Coiled-Coil Domain Containing 9B [46]
CCDC124 CCDC124 Coiled-Coil Domain Containing 124 [44]
CCDC85C CCDC85C Coiled-Coil Domain Containing 85C [44]
CCT3 CCT3 T-complex protein 1 subunit gamma [24]
CCT6A CCT6A T-complex protein 1 subunit zeta [24]
CDC20 CDC20 Cell Division Cycle 20 [46]
CDC37 CDC37 Cell Division Cycle 37 [54]
CDC5L CDC5L Cell division cycle 5-like protein [24]
CDC73 CDC73 Parafibromin [24]
CDK1 CDK1 Cyclin-dependent kinase 1 [24]
CDK2 CDK2 Cyclin Dependent Kinase 2 [71]
CDV3 CDV3 CDV3 Homolog [44]
CELF1 CUGBP1 CUGBP Elav-Like Family Member 1 [24][44][45][46][72]
CELF2 CUGBP2/BRUNOL3 CUGBP Elav-Like Family Member 2 [44]
CELF3 CUGBP3/BRUNOL1 CUGBP Elav-Like Family Member 3 [44]
CENPB CENPB Major centromere autoantigen B [24]
CENPF CENPF Centromere Protein F [46]
CEP78 CEP78/CRDHL Centrosomal Protein 78 [44]
CEP85 CEP85/CCDC21 Centrosomal Protein 78 [45]
CERKL Ceramide-Kinase Like Ceramide Kinase Like [73]
CFL1 Cofilin-1 Cofilin-1 [24]
CHCHD3 CHCHD3 Coiled-coil-helix-coiled-coil-helix domain-containing protein 3, mitochondrial [24]
CHORDC1 CHORDC1/CHP1 Cysteine and histidine-rich domain-containing protein 1 [24]
CIRBP CIRP Cold Inducible RNA Binding Protein [44][46][74]
CIT CIT Citron Rho-interacting kinase [24]
CLIC4 CLIC4 Chloride intracellular channel protein 4 [24]
CLNS1A CLNS1A Chloride Nucleotide-Sensitive Channel 1A [44]
CLPP CLPP Caseinolytic Mitochondrial Matrix Peptidase Proteolytic Subunit [44]
CNBP ZNF9 CCHC-Type Zinc Finger Nucleic Acid Binding Protein [46][75]
CNN3 CNN3 Calponin-3 [24]
CNOT1 CNOT1/CCR4 CCR4-Not Transcription Complex Subunit 1 [24][45] yes[45][76]
CNOT10 CNOT10 CCR4-Not Transcription Complex Subunit 10 [45] yes[45]
CNOT11 CNOT11 CCR4-Not Transcription Complex Subunit 11 [45] yes[45]
CNOT2 CNOT2 CCR4-Not Transcription Complex Subunit 2 [45] yes[45]
CNOT3 CNOT3 CCR4-Not Transcription Complex Subunit 3 [45] yes[45]
CNOT4 CNOT4 CCR4-Not Transcription Complex Subunit 4 [45] yes[45]
CNOT6 CNOT6 CCR4-Not Transcription Complex Subunit 6 [45] yes[45]
CNOT6L CNOT6L CCR4-Not Transcription Complex Subunit 6L [45] yes[45]
CNOT7 CNOT7 CCR4-Not Transcription Complex Subunit 7 [45] yes[45]
CNOT8 CNOT8 CCR4-Not Transcription Complex Subunit 8 [45] yes[45]
CNOT9 CNOT9 CCR4-Not Transcription Complex Subunit 9 [45]
CORO1B CORO1B Coronin-1B [24]
CPB2 Carboxypeptidase B2 Carboxypeptidase B2 [77]
CPEB1 CPEB Cytoplasmic Polyadenylation Element Binding Protein 1 [78]
CPEB4 CPEB4 Cytoplasmic Polyadenylation Element Binding Protein 4 [44][45][46] yes[45]
CPSF3 CPSF3 Cleavage and polyadenylation specificity factor subunit 3 [24]
CPSF6 CPSF6 Cleavage and polyadenylation specificity factor subunit 6 [24]
CPSF7 CPSF7 Cleavage and polyadenylation specificity factor subunit 7 [24]
CPVL CPVL Carboxypeptidase, Vitellogenic Like [45] yes[45]
CRKL CRKL CRK Like Proto-Oncogene, Adaptor Protein [44]
CROCC CROCC Ciliary Rootlet Coiled-Coil, Rootletin [44]
CRYAB CRYAB Alpha-crystallin B chain [24]
CRYBG1 CRYBG1 Crystallin Beta-Gamma Domain Containing 1 [46]
CSDE1 CSDE1 Cold shock domain-containing protein E1 [24][44][45][46][60]
CSE1L CSE1L/XPO2/Exportin-2 Exportin-2 [24]
CSNK2A1 Casein Kinase 2 alpha Casein Kinase 2 Alpha 1 [79]
CSTB Cystatin B Cystatin B [44]
CSTF1 CSTF1 Cleavage stimulation factor subunit 1 [24]
CTNNA2 CTNNA2 Catenin alpha-2 [24]
CTNND1 CTNND1 Catenin delta-1 [24]
CTTNBP2NL CTTNBP2NL CTTNBP2 N-terminal-like protein [24]
CWC22 CWC22 Pre-mRNA-splicing factor CWC22 homolog [24]
DAZAP1 DAZAP1 DAZ-associated protein 1 [24][44][45][46]
DAZAP2 PRTB DAZ Associated Protein 2 [80]
DAZL DAZL1 Deleted In Azoospermia Like [81]
DCD DCD Dermcidin [24]
DCP1A DCP1a Decapping mRNA 1a [24][44][78] yes[35]
DCP1B DCP1b Decapping mRNA 1b [44][46] yes[35]
DCP2 DCP2 Decapping mRNA 2 [45]
DCTN1 DCTN1 Dynactin subunit 1 [24]
DDX1 DEAD box protein 1 DEAD-Box Helicase 1 [24][44][45][46][82]
DDX11 DEAD box protein 11 DEAD-Box Helicase 11 [46]
DDX19A DDX19A ATP-dependent RNA helicase DDX19A [24][60]
DDX21 DDX21 Nucleolar RNA helicase 2 [24] yes[35]
DDX3 DEAD box protein 3 DEAD-Box Helicase 3 [24][83][84]
DDX3X DDX3X DEAD-Box Helicase 3, X-Linked [44][45][46][85][86][60]
DDX3Y DDX3Y DEAD-Box Helicase 3, Y-Linked [44]
DDX31 DDX31 DEAD-Box Helicase 31 [46]
DDX47 DDX47 Probable ATP-dependent RNA helicase DDX47 [24]
DDX50 DDX50 ATP-dependent RNA helicase DDX50 [24] yes[35]
DDX58 RIG-I DExD/H-Box Helicase 58 [87]
DDX6 DEAD box protein 6 DEAD-Box Helicase 6 [24][44][45][56][88][78][48][89][46] yes[35][45]
DERA DERA Deoxyribose-Phosphate Aldolase [90]
DGCR8 DGCR8 DGCR8 Microprocessor Complex Subunit [46]
DHX30 DHX30 Putative ATP-dependent RNA helicase DHX30 [24][44] yes[35]
DHX33 DHX33 DEAH-Box Helicase 33 [44]
DHX36 RHAU DEAH-Box Helicase 36 [44][45][25][46]
DHX57 DHX57 DExH-Box Helicase 57 [45][46]
DHX58 LGP2 DExH-Box Helicase 58 [87]
DIDO1 DIDO1 Death Inducer-Obliterator 1 [46]
DIS3L2 DIS3L2/FAM3A DIS3 Like 3'-5' Exoribonuclease 2 [44]
DISC1 Disrupted in Schizophrenia 1 Disrupted In Schizophrenia 1 [91]
DKC1 DKC1 dyskerin; H/ACA ribonucleoprotein complex subunit 4 [24][92]
DNAI1 Axonemal Dynein Intermediate Chain 1 Dynein Axonemal Intermediate Chain 1 [93]
DNAJA1 DNAJA1 DnaJ homolog subfamily A member 1 [24]
DNAJC8 DNAJC8 DnaJ homolog subfamily C member 8 [24]
DOCK4 DOCK4 Dedicator Of Cytokinesis 4 [46]
DPYSL2 DPYSL2 Dihydropyrimidinase-related protein 2 [24]
DPYSL3 DPYSL3 Dihydropyrimidinase-related protein 3 [24]
DROSHA DROSHA Drosha Ribonuclease III [44]
DSP DSP Desmoplakin [24][44]
DST DST Dystonin [24]
DSTN DSTN Destrin [24]
DTL DTL Denticleless E3 Ubiquitin Protein Ligase Homolog [46]
DTX3L DTX3L E3 ubiquitin-protein ligase DTX3L [24]
DUSP12 DUSP12/YVH1 Dual Specificity Phosphatase 12 [94]
DYNC1H1 Cytoplasmic Dynein Heavy Chain 1 Dynein Cytoplasmic 1 Heavy Chain 1 [93]
DYNLL1 Cytoplasmic Dynein Light Polypeptide Dynein Light Chain LC8-Type 1 [44][95]
DYNLL2 DYNLL2 Dynein light chain 2, cytoplasmic [24]
DYRK3 DYRK3 Dual Specificity Tyrosine Phosphorylation Regulated Kinase 3 [96]
DZIP1 DZIP1 DAZ Interacting Zinc Finger Protein 1 [97]
DZIP3 DZIP3 DAZ Interacting Zinc Finger Protein 3 [45]
EDC3 EDC3 Enhancer of mRNA Decapping 3 [44][45][46] yes[45]
EDC4 EDC4 Enhancer of mRNA-Decapping protein 4 [24][44][46] yes[35]
EIF1 EIF1 Eukaryotic Translation Initiation Factor 1 [44]
EIF2A EIF2A Eukaryotic Translation Initiation Factor 2A [36][24][52][98]
EIF2AK2 Protein Kinase R/PKR Eukaryotic Translation Initiation Factor 2 Alpha Kinase 2 [68][87][99]
EIF2B1-5 EIF2B Eukaryotic Translation Initiation Factor 2B [98]
EIF2S1 EIF2A subunit 1 Eukaryotic Translation Initiation Factor 2 Subunit Alpha [24]
EIF2S2 EIF2A subunit 2 Eukaryotic Translation Initiation Factor 2 Subunit Beta [24]
EIF3A EIF3A Eukaryotic Translation Initiation Factor 3 Subunit A [24][44][49][34][100][46]
EIF3B EIF3B Eukaryotic Translation Initiation Factor 3 Subunit B [36][24][80][101][102]
EIF3C EIF3C Eukaryotic Translation Initiation Factor 3 Subunit C [44]
EIF3D EIF3D Eukaryotic translation initiation factor 3 subunit D [24][44][60]
EIF3E EIF3E Eukaryotic translation initiation factor 3 subunit E [24][44][60]
EIF3F EIF3F Eukaryotic translation initiation factor 3 subunit F [24]
EIF3G EIF3G Eukaryotic translation initiation factor 3 subunit G [24][44][60][46]
EIF3H EIF3H Eukaryotic translation initiation factor 3 subunit H [24][44][46]
EIF3I EIF3I Eukaryotic translation initiation factor 3 subunit I [24][46]
EIF3J EIF3J Eukaryotic translation initiation factor 3 subunit J [24][44]
EIF3K EIF3K Eukaryotic translation initiation factor 3 subunit K [24]
EIF3L EIF3L Eukaryotic translation initiation factor 3 subunit L [24][44][60]
EIF3M EIF3M Eukaryotic translation initiation factor 3 subunit M [24]
EIF4A1 EIF4A1 Eukaryotic Translation Initiation Factor 4A1 [24][44][103][46]
EIF4A2 EIF4A2 Eukaryotic Translation Initiation Factor 4A2 [44][104][46]
EIF4A3 EIF4A3 Eukaryotic Translation Initiation Factor 4A3 [44]
EIF4B EIF4B Eukaryotic translation Initiation factor 4B [24][44][46]
EIF4E EIF4E Eukaryotic Translation Initiation Factor 4E [100][98][3][105][70][106][107][36] yes[36]
EIF4E2 EIF4E2 Eukaryotic Translation Initiation Factor 4E Family Member 2 [45][107] yes[45]
EIF4E3 EIF4E3 Eukaryotic Translation Initiation Factor 4E Family Member 3 [107]
EIF4ENIF1 EIF4ENIF1 Eukaryotic Translation Initiation Factor 4E Nuclear Import Factor 1 [44][45] yes[45]
EIF4G1 EIF4G1 Eukaryotic Translation Initiation Factor 4G1 [24][44][100][98][3][105][108][109][80][110][34][46]
EIF4G2 EIF4G2 Eukaryotic Translation Initiation Factor 4G2 [24][45]
EIF4G3 EIF4G3 Eukaryotic Translation Initiation Factor 4G3 [44]
EIF4H EIF4H Eukaryotic translation Initiation factor 4H [24][44][46]
EIF5A EIF5A Eukaryotic Translation Initiation Factor 5A [101]
ELAVL1 HuR ELAV Like RNA Binding Protein 1 [24][34][44][111][100][112][105][106][80][95][113][114][46] yes[35]
ELAVL2 ELAVL2 ELAV-like protein 2 [24][44] yes[35]
ELAVL3 ELAVL3/HuC ELAV Like RNA Binding Protein 3 [44]
ELAVL4 HuD ELAV Like RNA Binding Protein 4 [44][115]
ENC1 ENC1 Ectodermal-Neural Cortex 1 [46]
ENDOV EndoV Endonuclease V [116]
ENTPD1 ENTPD1 Ectonucleoside Triphosphate Diphosphohydrolase 1 [44]
EP400 EP400 E1A Binding Protein P400 [46]
EPPK1 EPPK1 Epiplakin [24]
ETF1 ETF1 Eukaryotic peptide chain release factor subunit 1 [24]
EWSR1 EWSR1 EWS RNA Binding Protein 1 [117][118][46]
FABP5 FABP5 Fatty Acid Binding Protein 5 [44]
FAM120A FAM120A/OSSA Constitutive coactivator of PPAR-gamma-like protein 1 [24][44][45] yes[35]
FAM120C FAM120C Family With Sequence Similarity 120C [44][45]
FAM168A FAM168A Family With Sequence Similarity 168 Member A [46]
FAM168B FAM168B/MANI Family With Sequence Similarity 168 Member B [44]
FAM83H FAM83H Family With Sequence Similarity 83 Member H [46]
FAM98A FAM98A Family With Sequence Similarity 98 Member A [24][44][119][46]
FAM98C FAM98C Family With Sequence Similarity 98 Member C [46]
FASTK FAST Fas Activated Serine/Threonine Kinase [36] yes[36]
FBL FBL rRNA 2-O-methyltransferase fibrillarin [24]
FBRSL1 Fibrosin Like 1 Fibrosin Like 1 [45]
FHL1 FHL1 Four and a half LIM domains protein 1 [24]
FKBP1A FKBP1A FKBP Prolyl Isomerase 1A [46]
FLNB FLNB Filamin-B [24]
FMR1 FMRP Fragile X Mental Retardation 1 [22][24][44][45][69][70][105][120][121][94][60][46]
FNDC3B FNDC3B Fibronectin type III domain-containing protein 3B [24][45][46]
FSCN1 FSCN1 Fascin [24]
FTSJ3 FTSJ3 pre-rRNA processing protein FTSJ3 [24]
FUBP1 FUBP1 Far Upstream Element Binding Protein 1 [44][46]
FUBP3 FUBP3 Far upstream element-binding protein 3 [24][44][45][46]
FUS FUS FUS RNA Binding Protein [24][44][49][117][118][122][123][124][125][126][127][128][46]
FXR1 FXR1 FMR1 Autosomal Homolog 1 [24][44][45][120][105][106][129][46]
FXR2 FXR2 FMR1 Autosomal Homolog 2 [24][44][45][120][105][46]
G3BP1 G3BP1 G3BP Stress Granule Assembly Factor 1 [24][44][45][67][99][68][130][131][36][106][132][129][133][60][46]
G3BP2 G3BP2 G3BP Stress Granule Assembly Factor 2 [24][44][45][134][135][60][46]
GABARAPL2 GABARAPL2/GEF2/ATG8 GABA Type A Receptor Associated Protein Like 2 [44]
GAK GAK Cyclin G Associated Kinase [46]
GAR1 GAR1 H/ACA Ribonucleoprotein Complex Subunit 1 [92]
GCA Grancalcin Grancalcin [44]
GEMIN5 Gemin-5 Gem Nuclear Organelle Associated Protein 5 [108]
GFPT1 GFPT1 Glutamine—fructose-6-phosphate aminotransferase [isomerizing] 1 [24]
GIGYF1 GIGYF1/PERQ1 GRB10 Interacting GYF Protein 1 [44]
GIGYF2 GIGYF2/TNRC15/PARK11/PERQ2 GRB10 Interacting GYF Protein 2 [44][45] yes[45]
GLE1 GLE1 GLE1, RNA Export Mediator [45][136][137]
GLO1 Glyoxalase Glyoxalase [44]
GLRX3 GLRX3/Glutaredoxin 3/TNLX2 Glutaredoxin 3 [44]
GLUD1 GLUD1 Glutamate Dehydrogenase 1 [46]
GNB2 GNB2 Guanine nucleotide-binding protein G(I)/G(S)/G(T) subunit beta-2 [24]
GOLGA2 Golgin A2 Golgin A2 [44]
GPAT3 GPAT3 Glycerol-3-Phosphate Acyltransferase 3 [46]
GRB2 GRB2/ASH Growth Factor Receptor Bound Protein 2 [44]
GRB7 GRB7 Growth Factor Receptor Bound Protein 7 [138][139]
GRSF1 GRSF1 G-Rich RNA Sequence Binding Factor 1 [44][45]
GSPT1 eRF3 G1 To S Phase Transition 1 [44][140]
GTF2I GTF2I General Transcription Factor IIi [46]
GTF3C1 GTF3C1 General Transcription Factor IIIC Subunit 1 [46]
GTF3C4 GTF3C4 General Transcription Factor IIIC Subunit 4 [46]
H1F0 H1F0 Histone H1.0 [24]
H1FX H1FX Histone H1x [24]
H2AFV H2AFV Histone H2A.V [24]
HABP4 Ki-1/57 Hyaluronan Binding Protein 4 [141]
HDAC6 HDAC6 Histone Deacetylase 6 [86][132][60]
HDLBP HDL-Binding Protein/VGL/Vigilin High Density Lipoprotein Binding Protein [44]
HELZ HELZ Probable helicase with zinc finger domain [24][44][45] yes[45]
HELZ2 HELZ2 Helicase with zinc finger domain 2 [24]
HMGA1 HMGA1 High mobility group protein HMG-I/HMG-Y [24]
HMGB3 HMGB3 High mobility group protein B3 [24]
HMGN1 HMGN1 Non-histone chromosomal protein HMG-14 [24]
HNRNPA1 HnRNPA1 Heterogeneous Nuclear Ribonucleoprotein A1 [24][44][49][142][143][144][145]
HNRNPA2B1 HnRNPA2/B1 Heterogeneous Nuclear Ribonucleoprotein A2/B1 [24][44][146][60]
HNRNPA3 HNRNPA3 Heterogeneous nuclear ribonucleoprotein A3 [24][44]
HNRNPAB HNRNPAB Heterogeneous nuclear ribonucleoprotein A/B [24][44][45]
HNRNPD HNRNPD Heterogeneous nuclear ribonucleoprotein D [44]
HNRNPDL HNRNPDL Heterogeneous nuclear ribonucleoprotein D-like [44]
HNRNPF HNRNPF Heterogeneous nuclear ribonucleoprotein F [44]
HNRNPH1 HNRNPH1 Heterogeneous nuclear ribonucleoprotein H1 [44]
HNRNPH2 HNRNPH2 Heterogeneous nuclear ribonucleoprotein H2 [24]
HNRNPH3 HNRNPH3 Heterogeneous nuclear ribonucleoprotein H3 [44]
HNRNPK HNRNPK Heterogeneous Nuclear Ribonucleoprotein K [24][114][147]
HNRNPUL1 HNRNPUL1 Heterogeneous nuclear ribonucleoprotein U-like protein 2 [24]
HSBP1 HSBP1 Heat Shock Factor Binding Protein 1 [44]
HSP90AA1 HSP90 Heat shock protein HSP 90-alpha [24]
HSPA4 HSP70 RY Heat shock 70 kDa protein 4 [24]
HSPA9 HSP70 9B Stress-70 protein, mitochondrial [24]
HSPB1 HSP27 Heat Shock Protein Family B (Small) Member 1 [24][148] yes[35]
HSPB8 HSPB8 Heat Shock Protein Family B (Small) Member 8 [149]
HSPBP1 HSPBP1 HSPA (Hsp70) Binding Protein 1 [150]
HSPD1 HSPD1 60 kDa heat shock protein, mitochondrial [24][44]
HTT Huntingtin Huntingtin [66]
IBTK IBTK Inhibitor Of Bruton Tyrosine Kinase [45]
IFIH1 MDA5 Interferon Induced With Helicase C Domain 1 [87]
IGF2BP1 IGF2BP1 Insulin-like Growth Factor 2 mRNA-binding protein 1 [24][44][45] yes[35]
IGF2BP2 IGF2BP2 Insulin-like Growth Factor 2 mRNA-binding protein 2 [24][44][45] yes[35]
IGF2BP3 IGF2BP3 Insulin-like Growth Factor 2 mRNA Binding Protein 3 [24][44][45][134] yes[35]
IK IK Protein Red [24]
ILF3 NF90 Interleukin Enhancer Binding Factor 3 [151] yes[35]
IPO7 IPO7 Importin-7 [24]
IPPK IP5K Inositol-Pentakisphosphate 2-Kinase [152]
ITGB1 ITGB1 Integrin beta-1 [24]
JMJD6 JMJD6 Arginine Demethylase and Lysine Hydroxylase [133]
KANK2 KANK2 KN motif and ankyrin repeat domain-containing protein 2 [24]
KEAP1 KEAP1/KLHL19 Kelch Like ECH Associated Protein 1 [44]
KHDRBS1 Sam68 KH RNA Binding Domain Containing, Signal Transduction Associated 1 [24][153][154][155]
KHDRBS3 KHDRBS3 KH domain-containing, RNA-binding, signal transduction-associated protein 3 [24]
KHSRP KSRP/FBP2 KH-Type Splicing Regulatory Protein [24][44][156]
KIAA0232 KIAA0232 KIAA0232 [45] yes[45]
KIAA1524 CIP2A Protein CIP2A [24]
KIF1B KIF1B Kinesin Family Member 1B [45]
KIF13B KIF13B/GAKIN Kinesin Family Member 13B [44]
KIF23 KIF23 Kinesin-like protein KIF23 [24] yes[35]
KIF2A Kinesin Heavy Chain Member 2 Kinesin Family Member 2A [93]
KLC1 Kinesin Light Chain 1 Kinesin Light Chain 1 [93]
KPNA1 Importin-ɑ5 Karyopherin Subunit Alpha 1 [24][44][157]
KPNA2 Importin-ɑ1 Karyopherin Subunit Alpha 2 [24][157][158][137]
KPNA3 Importin-ɑ4 Karyopherin Subunit Alpha 3 [44][157]
KPNA6 Importin-ɑ7 Importin subunit alpha [24]
KPNB1 Importin-β1 Karyopherin Subunit Beta 1 [24][157][137][60]
L1RE1 LINE1 ORF1p LINE1 ORF1 protein [24][49]
LANCL1 LanC Like 1 LanC Like 1 [44]
LARP1 LARP1 La-related protein 1 [24]
LARP1B LARP1B La-related protein 1b [45]
LARP4 La-Related protein 4 La Ribonucleoprotein Domain Family Member 4 [24][44][45][159]
LARP4B LARP4B La Ribonucleoprotein Domain Family Member 4B [44][45]
LASP1 LIM And SH3 Protein 1/MLN50 LIM And SH3 Protein 1 [44]
LBR LBR Lamin-B receptor [24]
LEMD3 LEMD3 Inner nuclear membrane protein Man1 [24]
LIG3 DNA Ligase 3 DNA Ligase 3 [44]
LIN28A LIN28A Lin-28 Homolog A [44][160]
LIN28B LIN28B Lin-28 Homolog B [44][160]
LMNA LMNA Prelamin-A/C [24]
LPP LPP Lipoma-preferred partner [24]
LSM1 LSM1 LSM1 Homolog, mRNA Degradation Associated [44] yes[161]
LSM12 LSM12 LSM12 Homolog [44][45]
LSM14A RAP55 LSM14A, mRNA Processing Body Assembly Factor [24][44][45][162][163] yes[35][45]
LSM14B LSM14B Protein LSM14 homolog B [24][44][45] yes[35]
LSM3 LSM3 U6 snRNA-associated Sm-like protein LSm3 [24] yes[161]
LUC7L LUC7L Putative RNA-binding protein Luc7-like 1 [24]
LUZP1 LUZP1 Leucine zipper protein 1 [24][45]
MACF1 MACF1 Microtubule-actin cross-linking factor 1, isoforms 1/2/3/5 [24][60]
MAEL MAEL Maelstrom Spermatogenic Transposon Silencer [164]
MAGEA4 MAGEA4 Melanoma-associated antigen 4 [24]
MAGED1 MAGED1 Melanoma-associated antigen D1 [24][44][45]
MAGED2 MAGED2 Melanoma-associated antigen D2 [24]
MAGOHB MAGOHB Protein mago nashi homolog 2 [24]
MAP1LC3A LC3-I Microtubule Associated Protein 1 Light Chain 3 Alpha [165][166]
MAP4 MAP4 Microtubule-associated protein 4 [24]
MAPK1IP1L MAPK1IP1L Mitogen-Activated Protein Kinase 1 Interacting Protein 1 Like [44]
MAP4K4 MAP4K4 Mitogen-activated protein kinase kinase kinase kinase 4 [24]
MAPK8 JNK1 Mitogen-Activated Protein Kinase 8 [167]
MAPRE1 MAPRE1 Microtubule-associated protein RP/EB family member 1 [24]
MAPRE2 MAPRE2 Microtubule Associated Protein RP/EB Family Member 2 [44]
MARF1 MARF1 Meiosis Regulator And mRNA Stability Factor 1 [45] yes[45]
MARS MARS Methionine—tRNA ligase, cytoplasmic [24]
MBNL1 MBNL1 Muscleblind Like Splicing Regulator 1 [82]
MBNL2 MBNL2 Muscleblind Like Splicing Regulator 2 [45]
MCM4 MCM4 DNA replication licensing factor MCM4 [24]
MCM5 MCM5 DNA replication licensing factor MCM5 [24]
MCM7 MCM7 DNA replication licensing factor MCM7 [24] yes[35]
METAP1 METAP1 Methionine aminopeptidase [24]
METAP2 METAP2 Methionyl Aminopeptidase 2 [44]
MCRIP1 FAM195B/GRAN2 Granulin-2 [44][45][89]
MCRIP2 FAM195A/GRAN1 Granulin-1 [45][89]
MEX3A MEX3A RNA-binding protein MEX3A [24] yes[35]
MEX3B MEX3B Mex-3 RNA Binding Family Member B [44][168]
MEX3C MEX3C Mex-3 RNA Binding Family Member C [44][169]
MEX3D MEX3D Mex-3 RNA Binding Family Member D [45]
MFAP1 MFAP1 Microfibrillar-associated protein 1 [24]
MKI67 MKI67 Antigen KI-67 [24]
MKRN2 MKRN2 Makorin Ring Finger Protein 2 [44][45]
MOV10 MOV-10 Mov10 RISC Complex RNA Helicase [24][45][48] yes[35][45]
MSH6 MSH6 DNA mismatch repair protein Msh6 [24]
MSI1 Musashi-1 Musashi RNA Binding Protein 1 [44][163] yes[35]
MSI2 MSI2 RNA-binding protein Musashi homolog 2 [24][44]
MTHFD1 MTHFD1 C-1-tetrahydrofolate synthase, cytoplasmic [24]
MTHFSD MTHFSD Methenyltetrahydrofolate Synthetase Domain Containing [170]
MTOR MTOR Mechanistic Target Of Rapamycin [96][171]
MYO6 MYO6 Unconventional myosin-VI [24]
NCOA3 SRC-3 Nuclear Receptor Coactivator 3 [172]
NDEL1 NUDEL/MITAP1/EOPA NudE Neurodevelopment Protein 1 Like 1 [44]
NELFE NELF-E/RD Negative Elongation Factor Complex Member E [44]
NEXN NEXN Nexilin [24]
NXF1 NXF1/MEX67/TAP Nuclear RNA Export Factor 1 [45][60]
NKRF NRF NFK-B Repressing Factor [44]
NOLC1 Nucleolar And Coiled-Body Phosphoprotein 1/NOPP140 Nucleolar And Coiled-Body Phosphoprotein 1 [44]
NONO NonO Non-POU Domain Containing Octamer Binding [24][173]
NOP58 NOP58 Nucleolar protein 58 [24] yes[35]
NOSIP NOSIP Nitric oxide synthase-interacting protein [24]
NOVA2 NOVA2 NOVA Alternative Splicing Regulator 2 [44]
NRG2 Neuregulin-2 Neuregulin-2 [102]
NSUN2 NSUN2 tRNA (cytosine(34)-C(5))-methyltransferase [24]
NTMT1 NTMT1 N-terminal Xaa-Pro-Lys N-methyltransferase 1 [24]
NUDC NUDC Nuclear migration protein nudC [24]
NUFIP1 NUFIP NUFIP1, FMR1 Interacting Protein 1 [105]
NUFIP2 NUFIP2 Nuclear fragile X mental retardation-interacting protein 2 [24][44][45][89][60]
NUPL2 NUPL2 Nucleoporin Like 2 [137]
NUP153 NUP153 Nucleoporin 153 [44]
NUP205 NUP205 Nuclear pore complex protein Nup205 [24][137]
NUP210 NUP210/GP210 Nucleoporin 210 [137]
NUP214 NUP214 Nucleoporin 214 [137]
NUP50 NUP50 Nucleoporin 50 [137]
NUP58 NUP58/NUPL1 Nucleoporin 58 [137]
NUP85 NUP85 Nucleoporin 85 [137]
NUP88 NUP88 Nucleoporin 88 [137]
NUP98 NUP98/NUP96 Nuclear pore complex protein Nup98-Nup96 [24][137][60]
OASL OASL/OASL1 2'-5'-Oligoadenylate Synthetase Like [174]
OAS1 OAS 2′–5′ oligoadenylate synthetase [87]
OAS2 OAS2 2'-5'-Oligoadenylate Synthetase 2 [99]
OGFOD1 TPA1 2-Oxoglutarate And Iron Dependent Oxygenase Domain Containing 1 [175]
OGG1 OGG1 8-Oxoguanine DNA Glycosylase [176]
OSBPL9 Oxysterol Binding Protein Like 9 Oxysterol Binding Protein Like 9 [44]
OTUD4 OTUD4/HIN1 OTU Deubiquitinase 4 [44][45][177]
P4HB Prolyl 4-Hydroxylase Subunit Beta Prolyl 4-Hydroxylase Subunit Beta [44]
PABPC1 PABP1 Poly(A) Binding Protein Cytoplasmic 1 [24][44][45][148][112][55][120][70][105][134]
PABPC4 PABPC4 Polyadenylate-binding protein 4 [24][44][45]
PAK4 PAK4 Serine/threonine-protein kinase PAK 4 [24][44]
PALLD Palladin Palladin [24]
PARG PARG/PARG99/PARG102 Poly(ADP-Ribose) Glycohydrolase [178]
PARK7 PARK7/DJ-1 Parkinsonism Associated Deglycase [179] yes[179]
PARN PARN/DAN Poly(A)-Specific Ribonuclease [44]
PARP12 PARP-12/ARTD12 Poly(ADP-Ribose) Polymerase Family Member 12 [45][178][180]
PARP14 PARP-14 Poly(ADP-Ribose) Polymerase Family Member 14 [178]
PARP15 PARP-15 Poly(ADP-Ribose) Polymerase Family Member 15 [178]
PATL1 PATL1 PAT1 Homolog 1, Processing Body mRNA Decay Factor [44][45] yes[45]
PAWR PAWR PRKC apoptosis WT1 regulator protein [24]
PCBP1 PCBP1/HNRNPE1 Poly(RC) Binding Protein 1 [44][45]
PCBP2 PCBP2/HNRNPE2 Poly(RC) Binding Protein 2 [24][44][45][77]
PCNA PCNA Proliferating cell nuclear antigen [24]
PDAP1 PDAP1 PDGFA Associated Protein 1 [44]
PDCD4 PDCD4 Programmed Cell Death 4 [181]
PDCD6IP PDCD6IP Programmed cell death 6-interacting protein [24]
PDIA3 PDIA3 Protein Disulfide Isomerase Family A Member 3 [44]
PDLIM1 PDLIM1 PDZ and LIM domain protein 1 [24]
PDLIM4 PDLIM4 PDZ and LIM domain protein 4 [24]
PDLIM5 PDLIM5 PDZ and LIM domain protein 5 [24]
PDS5B PDS5B Sister chromatid cohesion protein PDS5 homolog B [24]
PEF1 PEF1 Penta-EF-Hand Domain Containing 1 [44]
PEG10 PEG10 Paternally Expressed 10 [45]
PELO PELO Protein pelota homolog [24]
PEPD Peptidase D Peptidase D [44]
PEX11B PEX11B Peroxisomal Biogenesis Factor 11 Beta [44]
PFDN4 PFDN4 Prefoldin subunit 4 [24]
PFN1 Profilin 1 Profilin 1 [24][59]
PFN2 Profilin 2 Profilin 2 [24][59]
PGAM5 PGAM5 Serine/threonine-protein phosphatase PGAM5, mitochondrial [24]
PGP PGP/G3PP Phosphoglycolate Phosphatase [44]
PHB2 Prohibitin 2 Prohibitin 2 [21]
PHLDB2 PHLDB2 Pleckstrin homology-like domain family B member 2 [24]
PKP1 Plakophilin 1 Plakophilin 1 [129]
PKP2 Plakophilin 2 Plakophilin 2 [24]
PKP3 Plakophilin 3 Plakophilin 3 [129]
PNPT1 PNPase I Polyribonucleotide Nucleotidyltransferase 1 [44]
POLR2B POLR2B DNA-directed RNA polymerase [24][60]
POM121 POM121 POM121 Transmembrane Nucleoporin [137]
POP7 RPP20 POP7 Homolog, Ribonuclease P/MRP Subunit [131]
PPME1 PPME1 Protein phosphatase methylesterase 1 [24]
PPP1R8 PPP1R8 Protein Phosphatase 1 Regulatory Subunit 8 [44]
PPP1R10 PPP1R10 Serine/threonine-protein phosphatase 1 regulatory subunit 10 [24][60]
PPP1R18 PPP1R18 Phostensin [24]
PPP2R1A PPP2R1A Serine/threonine-protein phosphatase 2A 65 kDa regulatory subunit A alpha isoform [24][60]
PPP2R1B PPP2R1B Serine/threonine-protein phosphatase 2A 65 kDa regulatory subunit A beta isoform [44]
PQBP1 PQBP-1 Polyglutamine Binding Protein 1 [182]
PRDX1 PRDX1 Peroxiredoxin-1 [24][44]
PRDX6 PRDX6 Peroxiredoxin-6 [24]
PRKAA2 AMPK-a2 Protein Kinase AMP-Activated Catalytic Subunit Alpha 2 [20]
PRKCA PKC-ɑ Protein Kinase C Alpha [134]
PRKRA PACT Protein Activator Of Interferon Induced Protein Kinase EIF2AK2 [24][54]
PRMT1 PRMT1 Protein arginine N-methyltransferase 1 [24]
PRMT5 PRMT5 Protein arginine N-methyltransferase 5 [24]
PRRC2A PRRC2A Proline Rich Coiled-Coil 2A [24][44][45]
PRRC2B PRRC2B Proline Rich Coiled-Coil 2B [44][45]
PRRC2C PRRC2C Proline Rich Coiled-Coil 2C [24][44][45][60]
PSMD2 PSMD2 26S proteasome non-ATPase regulatory subunit 2 [24][183]
PSPC1 PSP1 Paraspeckle Component 1 [44]
PTBP1 PTBP1 Polypyrimidine tract-binding protein 1 [44]
PTBP3 PTBP3 Polypyrimidine tract-binding protein 3 [24][44][45]
PTGES3 PTGES3 Prostaglandin E synthase 3 [24]
PTK2 FAK Protein Tyrosine Kinase 2 [138]
PUM1 Pumilio-1 Pumilio homolog 1 [24][44][45] yes[35]
PUM2 Pumilio-2 Pumilio RNA Binding Family Member 2 [44][45][70]
PURA PURA Transcriptional activator protein Pur-alpha [24][44][124][126]
PURB PURB Transcriptional activator protein Pur-beta [24][44]
PWP1 PWP1 PWP1 Homolog, Endonuclein [44]
PXDNL PMR1 Peroxidasin Like [184]
PYCR1 PYCR1 Pyrroline-5-carboxylate reductase [24]
QKI QKI/HQK QKI, KH Domain Containing RNA Binding [44]
R3HDM1 R3HDM1 R3H Domain Containing 1 [44][45]
R3HDM2 R3HDM2 R3H Domain Containing 2 [45]
RAB1A RAB1A Ras-related protein Rab-1A [24][60]
RACGAP1 RACGAP1 Rac GTPase-activating protein 1 [24]
RACK1 RACK1 Receptor For Activated C Kinase 1 [21][110][185]
RAD21 RAD21 Double-strand-break repair protein rad21 homolog [24]
RAE1 RAE1 Ribonucleic Acid Export 1 [137]
RAN RAN RAN, Member RAS Oncogene Family [158][137]
RANBP1 RANBP1 Ran-specific GTPase-activating protein [24]
RANBP2 RANBP2/NUP358 RAN Binding Protein 2 [137]
RBBP4 RBBP4 Histone-binding protein RBBP4 [24]
RBFOX1 RBFOX1 RNA binding protein fox-1 homolog [24][186][187] yes[187]
RBFOX2 RBFOX2 RNA binding protein fox-1 homolog 2 [186]
RBFOX3 RBFOX3 RNA binding protein fox-1 homolog 3 [186]
RBM12B RBM12B RNA-binding protein 12B [24]
RBM15 RBM15 RNA-binding protein 15 [44]
RBM17 RBM17 RNA-binding protein 17 [44]
RBM25 RBM25 RNA-binding protein 25 [44]
RBM26 RBM26 RNA-binding protein 26 [24]
RBM3 RBM3 RNA-binding protein 3 [44]
RBM38 RBM38 RNA-binding protein 38 [44]
RBM4 RBM4 RNA Binding Motif Protein 4 [44][188]
RBM4B RBM4B RNA Binding Motif Protein 4B [44]
RBM42 RBM42 RNA Binding Motif Protein 42 [147]
RBM45 RBM45 RNA Binding Motif Protein 45 [189][190]
RBM47 RBM47 RNA Binding Motif Protein 47 [45]
RBMS1 RBMS1 RNA-binding motif, single-stranded-interacting protein 1 [24][44][45]
RBMS2 RBMS2 RNA-binding motif, single-stranded-interacting protein 2 [24][44][45]
RBMX RBMX RNA Binding Motif Protein, X-Linked [45]
RBPMS RBPMS RNA-binding protein with multiple splicing [191]
RC3H1 Roquin-1 Ring Finger And CCCH-Type Domains 1 [44][45][192]
RC3H2 MNAB Ring Finger And CCCH-Type Domains 2 [45][192]
RCC1 RCC1 Regulator of chromosome condensation [24]
RCC2 RCC2 Protein RCC2 [24]
RECQL RECQL1 RecQ Like Helicase [44]
RFC3 RFC3 Replication factor C subunit 3 [24]
RFC4 RFC4 Replication factor C subunit 4 [24]
RGPD3 RGPD3 RanBP2-like and GRIP domain-containing protein 3 [24]
RHOA RhoA Ras Homolog Family Member A [22]
RNASEL RNAse L Ribonuclease L [87][68]
RNF214 RNF214 RING finger protein 214 [24][44]
RNF219 RNF219 RING finger protein 219 [45] yes[45]
RNF25 RNF25 Ring Finger Protein 25 [44]
RNH1 RNH1 Ribonuclease inhibitor [24][53]
ROCK1 ROCK1 Rho Associated Coiled-Coil Containing Protein Kinase 1 [22]
RPS19 Ribosomal Protein S19 Ribosomal Protein S19 [100]
RPS3 40S Ribosomal Protein S3 40S Ribosomal Protein S3 [98][100] yes[35]
RPS6 Ribosomal Protein S6 Ribosomal Protein S6 [67][98][3][105][171]
RPS11 Ribosomal Protein S11 Ribosomal Protein S11 [44]
RPS24 Ribosomal Protein S24 Ribosomal Protein S24 [44]
RPS6KA3 RSK2 Ribosomal Protein S6 Kinase A3 [193]
RPS6KB1 S6K1 Ribosomal Protein S6 Kinase B1 [171]
RPS6KB2 S6K2 Ribosomal Protein S6 Kinase B2 [171]
RPTOR RAPTOR Regulatory Associated Protein of mTOR Complex 1 [88][96][171]
RSL1D1 RSL1D1 Ribosomal L1 domain-containing protein 1 [24]
RTCB RTCB tRNA-splicing ligase RtcB homolog, formerly C22orf28 [24][44]
RTRAF RTRAF (formerly C14orf166) RNA Transcription, Translation And Transport Factor [44]
S100A7A S100A7A Protein S100-A7A [24]
S100A9 S100A9 Protein S100-A9 [24] yes[35]
SAFB2 SAFB2 Scaffold attachment factor B2 [24][44] yes[35]
SAMD4A SMAUG1 Sterile Alpha Motif Domain Containing 4A [194]
SAMD4B SMAUG2 Sterile Alpha Motif Domain Containing 4B [44]
SCAPER SCAPER S-Phase Cyclin A Associated Protein In The ER [45]
SEC24C SEC24C Protein transport protein Sec24C [24][44]
SECISBP2 SECIS Binding Protein 2 SECIS Binding Protein 2 [44][45]
SERBP1 PAI-RBP1/SERBP1 SERPINE1 mRNA Binding Protein 1 [49][195][84]
SERPINE1 PAI-1/Serpin E1 Serpine Family E Member 1 [196]
SF1 SF1 Splicing Factor 1 [44]
SFN SFN 14-3-3 protein sigma [24]
SFPQ PSF Splicing Factor Proline And Glutamine Rich [24][173]
SFRS3 SFRS3 Serine/arginine-rich splicing factor 3 [24]
SIPA1L1 SIPA1L1 Signal-induced proliferation-associated 1-like protein 1 [24]
SIRT6 Sirtuin 6 Sirtuin 6 [197]
SLBP Stem-Loop Binding Protein Stem-Loop Binding Protein [44]
SMAP2 SMAP2 Small ArfGAP2 [45]
SMARCA1 SMARCA1/SNF2L1 Probable global transcription activator SNF2L1 [24]
SMC4 SMC4 Structural maintenance of chromosomes protein [24]
SMG1 SMG-1 SMG1, Nonsense Mediated mRNA Decay Associated PI3K Related Kinase [194][198]
SMG6 SMG6 SMG6, Nonsense Mediated mRNA Decay Factor [45]
SMG7 SMG7 SMG7, Nonsense Mediated mRNA Decay Factor [45] yes[45]
SMN1 Survival of Motor Neuron Survival Of Motor Neuron 1, Telomeric [131][199][200]
SMU1 SMU1 WD40 repeat-containing protein SMU1 [24]
SMYD5 SMYD5 SMYD Family Member 5 [44]
SND1 Tudor-SN Staphylococcal Nuclease And Tudor Domain Containing 1 [44][45][47][201]
SNRPF SNRPF Small nuclear ribonucleoprotein F [24]
SNTB2 SNTB2 Beta-2-syntrophin [24]
SOGA3 SOGA3 SOGA Family Member 3 [44]
SORBS1 SORBS1 Sorbin and SH3 domain-containing protein 1 [24]
SORBS3 Vinexin Sorbin And SH3 Domain Containing 3 [202]
SOX3 SOX3 SRY-Box 3 [44]
SPAG5 Astrin Sperm Associated Antigen 5 [88][171]
SPATS2 SPATS2/SPATA10/SCR59 Spermatogenesis Associated Serine Rich 2 [44]
SPATS2L SGNP Spermatogenesis Associated Serine Rich 2 Like [24][203]
SPECC1L SPECC1L Cytospin-A [24]
SQSTM1 SQSTM1/p62 Sequestosome 1 [63]
SRI SRI Sorcin [24][44]
SRP68 Signal Recognition Particle 68 Signal Recognition Particle 68 [44][48]
SRP9 SRP9 Signal Recognition Particle 9 [204]
SRRT SRRT Serrate RNA effector molecule homolog [24]
SRSF1 ASF/SF2 Serine And Arginine Rich Splicing Factor 1 [44][205]
SRSF3 SRp20 Serine And Arginine Rich Splicing Factor 3 [206][207][208][60]
SRSF4 SRSF4 Serine/arginine-rich splicing factor 4 [24]
SRSF5 SRSF5/SRP40 Serine/arginine-rich splicing factor 5 [44]
SRSF7 9G8 Serine And Arginine Rich Splicing Factor 7 [49]
SRSF9 SRSF9/SRP30C Serine/arginine-rich splicing factor 9 [44]
SS18L1 SS18L1/CREST SS18L1, nBAF Chromatin Remodeling Complex Subunit [209]
ST7 ST7/FAM4A1/HELG/RAY1/TSG7 Suppression Of Tumorigenicity 7 [45] yes[45]
STAT1 STAT1 Signal transducer and activator of transcription 1-alpha/beta [24]
STAU1 Staufen 1 Staufen Double-Stranded RNA Binding Protein 1 [24][44][112][70][210]
STAU2 Staufen 2 Staufen Double-Stranded RNA Binding Protein 2 [24][44][45][112] yes[35]
STIP1 STIP1/HOP Stress-induced-phosphoprotein 1 [24][54]
STRAP STRAP Serine-threonine kinase receptor-associated protein [24][44]
SUGP2 SUGP2 SURP and G-patch domain-containing protein 2 [24]
SUGT1 SUGT1 SGT1 Homolog, MIS12 Kinetochore Complex Assembly Cochaperone [45]
SUN1 SUN1 SUN domain-containing protein 1 [24]
SYCP3 SYCP3 Synaptonemal complex protein 3 [24]
SYK SYK Spleen Associated Tyrosine Kinase [139]
SYNCRIP SYNCRIP Heterogeneous nuclear ribonucleoprotein Q [24][44][45][211] yes[35]
TAGLN3 Transgelin 3 Transgelin 3 [44]
TAF15 TAF15 TATA-Box Binding Protein Associated Factor 15 [24][44][117][118][122][60]
TARDBP TDP-43 TAR DNA Binding Protein [24][113][212][213][143][146][103][190][214][215]
TBRG1 TBRG1 Transforming Growth Factor Beta Regulator 1 [44]
TCEA1 TCEA1 Transcription elongation factor A protein 1 [24]
TCP1 TCP1 T-complex protein 1 subunit alpha [24]
TDRD3 Tudor Domain Containing 3 Tudor Domain Containing 3 [44][45][84][216][217][218]
TDRD7 Tudor Domain Containing 7 Tudor Domain Containing 7 [45]
TERT TERT Telomerase Reverse Transcriptase [219]
THOC2 THOC2 THO Complex 2 [137]
THRAP3 THRAP3 Thyroid Hormone Receptor Associated Protein 3 [44]
TIA1 TIA-1 TIA1 Cytotoxic Granule Associated RNA Binding Protein [3][24][44][49][56][34][70][80][95][121][132][142][148][199][214][220][60]
TIAL1 TIAR TIA1 Cytotoxic Granule Associated RNA Binding Protein Like 1 [24][44][45][70][105][112][113][148][189][199][209]
TMEM131 TMEM131 Transmembrane Protein 131 [45] yes[45]
TMOD3 TMOD3 Tropomodulin-3 [24]
TNKS PARP-5a Tankyrase [178]
TNKS1BP1 TNKS1BP1 182 kDa tankyrase-1-binding protein [24][45] yes[45]
TNPO1 Transportin-1 Transportin-1/Karyopherin (Importin) Beta 2 [24][44][137][221][222]
TNPO2 Transportin-2 Transportin-2 [24][45]
TNRC6A TNRC6A Trinucleotide repeat-containing gene 6A protein [44][45] yes[45]
TNRC6B TNRC6B Trinucleotide repeat-containing gene 6B protein [24][44][45] yes[45]
TNRC6C TNRC6C Trinucleotide repeat-containing gene 6C protein [44][45] yes[45]
TOMM34 TOMM34 Mitochondrial import receptor subunit TOM34 [24]
TOP3B Topoisomerase (DNA) III Beta Topoisomerase (DNA) III Beta [45][217][223]
TPM1 TPM1 Tropomyosin alpha-1 chain [24]
TPM2 TPM2 Tropomyosin beta chain [24]
TPR TPR Translocated Promoter Region, Nuclear Basket Protein [137]
TRA2B TRA2B Transformer 2 Beta Homolog [45]
TRAF2 TRAF2 TNF Receptor Associated Factor 2 [109]
TRDMT1 DNMT2 tRNA Aspartic Acid Methyltransferase 1 [224]
TRIM21 TRIM21 E3 ubiquitin-protein ligase TRIM21 [24]
TRIM25 TRIM25 E3 ubiquitin/ISG15 ligase TRIM25 [24][44][60]
TRIM56 TRIM56 E3 ubiquitin-protein ligase TRIM56 [24][45][60]
TRIM71 TRIM71 E3 ubiquitin-protein ligase TRIM71 [44]
TRIP6 TRIP6 Thyroid receptor-interacting protein 6 [24][44]
TROVE2 RORNP TROVE Domain Family Member 2 [44]
TTC17 TTC17 Tetratricopeptide Repeat Domain 17 [45] yes[45]
TUBA1C TUBA1C Tubulin alpha-1C chain [24]
TUBA3C TUBA3C Tubulin alpha-3C/D chain [24]
TUBA4A TUBA4A Tubulin alpha-4A chain [24]
TUBB3 TUBB3 Tubulin beta-3 chain [24]
TUBB8 TUBB8 Tubulin beta-8 chain [24]
TUFM TUFM Elongation factor Tu, mitochondrial [24]
TXN TXN Thioredoxin [24]
TXNDC17 TXNDC17 Thioredoxin Domain Containing 17 [44]
U2AF1 U2AF1 Splicing factor U2AF 35 kDa subunit [24]
UBA1 UBA1 Ubiquitin-like modifier-activating enzyme 1 [24]
UBAP2 UBAP2 Ubiquitin-associated protein 2 [24][44][45][60]
UBAP2L UBAP2L Ubiquitin-associated protein 2-like [24][44][45][225][226][60]
UBB Ubiquitin Ubiquitin [114][132]
UBL5 Ubiquitin Like 5 Ubiquitin Like 5 [44]
UBQLN2 Ubiquilin 2 Ubiquilin 2 [227]
ULK1 ULK1 Unc-51 Like Autophagy Activating Kinase 1 [228]
ULK2 ULK2 Unc-51 Like Autophagy Activating Kinase 2 [228]
UPF1 UPF1 UPF1, RNA Helicase and ATPase [24][44][45][198][60] yes[35]
UPF2 UPF2 UPF2, RNA Helicase and ATPase [198]
UPF3B UPF3B UPF3B, Regulator of Nonsense Mediated mRNA Decay [44]
USP10 USP10 Ubiquitin Specific Peptidase 10 [24][44][45][67][34][185][60]
USP11 USP11 Ubiquitin Specific Peptidase 11 [44]
USP13 USP13 Ubiquitin Specific Peptidase 13 [229]
USP5 USP5 Ubiquitin carboxyl-terminal hydrolase 5 [24][229]
USP9X USP9X Ubiquitin Specific Peptidase 9, X-Linked [218]
UTP18 UTP18 UTP18, Small Subunit Processome Component [44]
VASP VASP Vasodilator-stimulated phosphoprotein [24]
VBP1 VBP1 VHL Binding Protein 1 [44]
VCP VCP Valosin Containing Protein [24][230][183][228]
WBP2 WBP2 WW Domain Binding Protein 2 [44]
WDR47 WDR47 WD Repeat Domain 47 [44]
WDR62 WDR62 WD Repeat Domain 62 [167]
XPO1 XPO1/CRM1 Exportin 1 [137]
XRN1 XRN1 5'-3' Exoribonuclease 1 [36][44][45] yes[36][45]
XRN2 XRN2 5'-3' Exoribonuclease 2 [44]
YARS YARS Tyrosine—tRNA ligase, cytoplasmic [24]
YBX1 YB-1 Y-Box Binding Protein 1 [24][44][49][48][82][94][231]
YBX3 YBX3/ZONAB Y-box-binding protein 3 [24][44][45]
YES1 YES1 Tyrosine-protein kinase Yes [24]
YLPM1 YLPM1 YLP Motif Containing 1 [44]
YTHDF1 YTHDF1 YTH domain family protein 1 [24][44][45][232][233]
YTHDF2 YTHDF2 YTH domain family protein 2 [24][44][45][232][233] yes[232][233]
YTHDF3 YTHDF3 YTH domain family protein 3 [24][234][44][45][232][233]
YWHAB 14-3-3 Tyrosine 3-Monooxygenase/Tryptophan 5-Monooxygenase Activation Protein Beta [24][168]
YWHAH 14-3-3 14-3-3 protein eta [24]
YWHAQ 14-3-3 14-3-3 protein theta [24]
ZBP1 ZBP1 Z-DNA Binding Protein 1 [235][236]
ZCCHC11 ZCCHC11 Zinc finger CCCH domain-containing protein 11 [45]
ZCCHC14 ZCCHC14 Zinc finger CCCH domain-containing protein 14 [45]
ZC3H11A ZC3H11A Zinc finger CCCH domain-containing protein 11a [44]
ZC3H14 ZC3H14 Zinc finger CCCH domain-containing protein 14 [24]
ZCCHC2 ZCCHC2 Zinc finger CCCH domain-containing protein 2 [45]
ZCCHC3 ZCCHC3 Zinc finger CCCH domain-containing protein 3 [45]
ZC3H7A ZC3H7A Zinc finger CCCH domain-containing protein 7A [24]
ZC3H7B ZC3H7B Zinc finger CCCH domain-containing protein 7B [24][44]
ZC3HAV1 PARP-13.1/PARP-13.2/ARTD13 Zinc Finger CCCH-Type Containing, Antiviral 1 [24][45][178] yes[35]
ZFAND1 ZFAND1 Zinc Finger AN1-Type Containing 1 [183]
ZFP36 TTP/TIS11 ZFP36 Ring Finger Protein/Trisetrapolin [36][44][167][237][238][239] yes[36]
ZNF598 ZNF598 Zinc finger protein 598 [45]
ZNF638 ZNF638 Zinc finger protein 638 [24]

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