Stress granule
Stress granules are dense aggregations in the cytosol composed of proteins & RNAs that appear when the cell is under stress. The RNA molecules stored are stalled translation pre-initiation complexes: failed attempts to make protein from mRNA. Stress granules are 100–200 nm in size, not surrounded by membrane, and associated with the endoplasmatic reticulum. Note that there are also nuclear stress granules. This article is about the cytosolic variety.
Proposed functions
The function of stress granules remains largely unknown. Stress granules have long been proposed to have a function to protect RNAs from harmful conditions, thus their appearance under stress. The accumulation of RNAs 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 mRNAs. Molecules can go down one of three paths: further storage, degradation, or re-initiation of translation. 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 mRNAs in transit between a state of storage and a state of degradation.
Efforts to identify all RNAs within stress granules in an unbiased way by sequencing RNA from biochemically purified stress granule "cores" have shown that RNAs are not recruited to stress granules in a sequence-specific manner, but rather generically, with longer and/or less-optimally translated transcripts being enriched. These data imply that the stress granule transcriptome is influenced the valency of RNA and by the rates of RNA run-off from polysomes. The latter is further supported by recent single molecule imaging studies. Furthermore, it was estimated that only about 15% of the total mRNA in the cell is localized to stress granules, suggesting that stress granules only influence a minority of mRNAs in the cell and may not be as important for mRNA processing as previously thought. That said, these studies represent only a snapshot in time, and it is likely that a larger fraction of mRNAs are at one point stored in stress granules due to those RNAs 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. Therefore, any association of mRNAs with stress granules may simply be a side effect of the association of partially unfolded RNA-binding proteins with stress granules, similar to the association of mRNAs with proteasomes.
Formation
Environmental stressors trigger cellular signaling which eventually leads to the formation of stress granules. In vitro, these stressors can include heat, cold, oxidative stress, endoplasmic reticulum stress, proteasome inhibition, hyperosmotic stress, ultraviolet radiation, inhibition of eIF4A, nitric oxide accumulation after treatment with 3-morpholinosydnonimine, perturbation of pre-mRNA splicing, and other stressors like puromycin that result in disassembled polysomes. Many of these stressors result in the activation of particular stress-associated kinases, translational inhibition and stress granule formation.Stress granule formation is often downstream of the stress-activated phosphorylation of eukaryotic translation initiation factor eIF2α, but this isn't true for all types of stressors that induce stress granules, 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. It has also been proposed that microtubules play a role in the formation of stress granules, maybe 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. Furthermore, many signaling molecules were shown to regulate the formation or dynamics of stress granules; these include the master energy sensor AMP-activated protein kinase, the O-GlcNAc transferase enzyme, and the pro-apoptotic kinase ROCK1.
Potential roles of RNA-RNA interactions
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. RNA-seq analyses demonstrate that these assemblies share a largely overlapping transcriptome with stress granules, with RNA enrichment in both being predominately based on the length of the RNA. Furthermore, stress granules contain many RNA helicases, including the DEAD/H-box helicases /DDX3, eIF4A1, and RHAU. In yeast, catalytic ded1 mutant alleles give rise to constitutive stress granules ATPase-deficient DDX3X mutant alleles are found in pediatric medulloblastoma, and these coincide with constitutive granular assemblies in patient cells. These mutant DDX3 proteins promote stress granule assembly in HeLa cells. In mammalian cells, RHAU mutants lead to reduced stress granule dynamics. 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. There is also evidence that RNA within stress granules is more compacted compared to RNA in the cytoplasm and that the RNA is preferentially post-translationally modified by N6-methyladenosine on its 5' ends. 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.Connection with processing bodies
Stress granules and 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. 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 mRNAs.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. While both stress granules and processing bodies are associated with mRNAs, processing bodies have been long proposed to be sites of mRNA degradation because they contain enzymes like DCP1/2 and XRN1 that are known to degrade mRNAs. However, others have demonstrated that mRNAs associated with processing bodies are largely translationally repressed but not degraded. It has also been proposed that mRNAs selected for degradation are passed from stress granules to processing bodies, though there is also data suggesting that processing bodies precede and promote stress granule formation.
Protein composition of stress granules
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. Importantly, different stressors can result in stress granules with different protein components. 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 and/or by fixing cells and using antibodies to detect the protein of interest along with known protein markers of stress granules.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.
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 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. 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.
Another proximity labeling method used to determine the proteome of stress granules is BioID. BioID is similar to the APEX approach, in that a biotinylating protein 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.
A curated database of stress granule-associated proteins can be found here.
The following is a list of proteins that have been demonstrated to localize to stress granules :
| Gene ID | Protein Name | Description | References | Also found in processing bodies? |
| ABCF1 | ABCF1 | ATP Binding Cassette Subfamily F Member 1 | ||
| ABRACL | ABRACL | ABRA C-Terminal Like | ||
| ACAP1 | ACAP1 | ArfGAP With Coiled-Coil, Ankyrin Repeat And PH Domains 1 | ||
| ACBD5 | ACBD5 | Acyl-CoA Binding Domain Containing 5 | ||
| ACTBL2 | ACTBL2 | Beta-actin-like protein 2 | yes | |
| ACTR1A | ACTR1A | Alpha-centractin | ||
| ACTR1B | ACTR1B | Beta-centractin | ||
| ADAR | ADAR1 | Adenosine Deaminase, RNA Specific | ||
| ADD1 | Adducin 1 | Adducin 1 | ||
| AGO1 | Argonaute 1/EIF2C1 | Argonaute 1, RISC Catalytic Component | yes | |
| AGO2 | Argonaute 2 | Argonaute 2, RISC Catalytic Component | yes | |
| AKAP9 | AKAP350 | A-Kinase Anchoring Protein 9 | ||
| AKAP13 | AKAP13/LBC | A-Kinase Anchoring Protein 13 | ||
| ALDH18A1 | ALDH18A1 | Delta-1-pyrroline-5-carboxylate synthase | ||
| ALG13 | ALG13 | ALG13, UDP-N-Acetylglucosaminyltransferase Subunit | ||
| ANKHD1 | ANKHD1 | Ankyrin Repeat and KH Domain Containing 1 | yes | |
| ANKRD17 | ANKRD17/MASK2/GTAR | Ankyrin Repeat Domain 17 | yes | |
| ANG | Angiogenin | Angiogenin | ||
| ANP32E | ANP32E | Acidic leucine-rich nuclear phosphoprotein 32 family member E | ||
| ANXA1 | ANXA1 | Annexin A1 | ||
| ANXA11 | ANXA11 | Annexin 11 | ||
| ANXA6 | ANXA6 | Annexin 6 | ||
| ANXA7 | ANXA7 | Annexin 7 | ||
| APEX1 | APEX1 | DNA- lyase | ||
| APOBEC3C | APOBEC3C | Apolipoprotein B mRNA Editing Enzyme Catalytic Subunit 3C | ||
| APOBEC3G | APOBEC3G | Apolipoprotein B mRNA Editing Enzyme Catalytic Subunit 3G | ||
| ARPC1B | ARPC1B | Actin-related protein 2/3 complex subunit 1B | ||
| AHSA1 | AHA1 | Activator Of HSP90 ATPase Activity 1 | ||
| AQR | AQR/IBP160 | Aquarius Intron-Binding Spliceosomal Factor | ||
| ARMC6 | ARMC6 | Armadillo Repeat Containing 6 | ||
| ASCC1 | ASCC1 | Activating Signal Cointegrator 1 Complex Subunit 1 | ||
| ASCC3 | ASCC3 | Activating Signal Cointegrator 1 Complex Subunit 3 | ||
| ATAD2 | ATAD2 | ATPase family AAA domain-containing protein 2 | ||
| ATAD3A | ATAD3A | ATPase family AAA domain-containing protein 3A | yes | |
| ATG3 | ATG3 | Autophagy Related 3 | ||
| ATP5A1 | ATP5A1 | ATP synthase subunit alpha, mitochondrial | ||
| ATP6V1G1 | ATP6V1G1/ATP6G | ATPase H+ Transporting V1 Subunit G1 | ||
| ATXN2 | Ataxin 2 | Ataxin 2 | ||
| ATXN2L | Ataxin-2 like | Ataxin 2 Like | ||
| BAG3 | BAG3 | BAG family molecular chaperone regulator 3 | ||
| BANF1 | BANF1 | Barrier-to-autointegration factor | ||
| BCCIP | BCCIP | BRCA2 And CDKN1A Interacting Protein | ||
| BCLAF1 | BCLAF1 | BCL2 Associated Transcription Factor 1 | ||
| BICC1 | BICC1 | BicC Family RNA Binding Protein 1 | ||
| BOLL | BOULE | Boule Homolog, RNA Binding Protein | ||
| BRAT1 | BRAT1 | BRCA1-associated ATM activator 1 | ||
| BRF1 | BRF1 | BRF1, RNA Polymerase III Transcription Initiation Factor Subunit | ||
| BTG3 | BTG3 | BTG Anti-Proliferation Factor 3 | yes | |
| C9orf72 | C9orf72 | Uncharacterized protein C9orf72 | ||
| C15orf52 | C15orf52 | Uncharacterized protein C15orf52 | ||
| C20orf27 | C20orf72 | Chromosome 20 Open Reading Frame 27 | ||
| C2CD3 | C2CD3 | C2 Calcium Dependent Domain Containing 3 | ||
| CALML5 | CALML5 | Calmodulin-like protein 5 | ||
| CALR | Calreticulin/CRT | Calreticulin | ||
| CAP1 | CAP1 | Adenylyl cyclase-associated protein 1 | ||
| CAPRIN1 | Caprin-1 | Cell Cycle Associated Protein 1 | ||
| CAPZA2 | CAPZA2 | F-actin-capping protein subunit alpha-2 | ||
| CARHSP1 | CARHSP1 | Calcium-regulated heat stable protein 1 | ||
| CASC3 | MLN51/BTZ | Cancer Susceptibility 3 | ||
| CBFB | CBFB | Core-binding factor subunit beta | ||
| CBX1 | CBX1 | Chromobox protein homolog 1 | ||
| CCAR1 | CARP-1 | Cell Division Cycle And Apoptosis Regulator 1 | ||
| CCDC124 | CCDC124 | Coiled-Coil Domain Containing 124 | ||
| CCDC85C | CCDC85C | Coiled-Coil Domain Containing 85C | ||
| CCT3 | CCT3 | T-complex protein 1 subunit gamma | ||
| CCT6A | CCT6A | T-complex protein 1 subunit zeta | ||
| CDC37 | CDC37 | Cell Division Cycle 37 | ||
| CDC5L | CDC5L | Cell division cycle 5-like protein | ||
| CDC73 | CDC73 | Parafibromin | ||
| CDK1 | CDK1 | Cyclin-dependent kinase 1 | ||
| CDK2 | CDK2 | Cyclin Dependent Kinase 2 | ||
| CDV3 | CDV3 | CDV3 Homolog | ||
| CELF1 | CUGBP1 | CUGBP Elav-Like Family Member 1 | ||
| CELF2 | CUGBP2/BRUNOL3 | CUGBP Elav-Like Family Member 2 | ||
| CELF3 | CUGBP3/BRUNOL1 | CUGBP Elav-Like Family Member 3 | ||
| CENPB | CENPB | Major centromere autoantigen B | ||
| CEP78 | CEP78/CRDHL | Centrosomal Protein 78 | ||
| CEP85 | CEP85/CCDC21 | Centrosomal Protein 78 | ||
| CERKL | Ceramide-Kinase Like | Ceramide Kinase Like | ||
| CFL1 | Cofilin-1 | Cofilin-1 | ||
| CHCHD3 | CHCHD3 | Coiled-coil-helix-coiled-coil-helix domain-containing protein 3, mitochondrial | ||
| CHORDC1 | CHORDC1/CHP1 | Cysteine and histidine-rich domain-containing protein 1 | ||
| CIRBP | CIRP | Cold Inducible RNA Binding Protein | ||
| CIT | CIT | Citron Rho-interacting kinase | ||
| CLIC4 | CLIC4 | Chloride intracellular channel protein 4 | ||
| CLNS1A | CLNS1A | Chloride Nucleotide-Sensitive Channel 1A | ||
| CLPP | CLPP | Caseinolytic Mitochondrial Matrix Peptidase Proteolytic Subunit | ||
| CNBP | ZNF9 | CCHC-Type Zinc Finger Nucleic Acid Binding Protein | ||
| CNN3 | CNN3 | Calponin-3 | ||
| CNOT1 | CNOT1/CCR4 | CCR4-Not Transcription Complex Subunit 1 | yes | |
| CNOT10 | CNOT10 | CCR4-Not Transcription Complex Subunit 10 | yes | |
| CNOT11 | CNOT11 | CCR4-Not Transcription Complex Subunit 11 | yes | |
| CNOT2 | CNOT2 | CCR4-Not Transcription Complex Subunit 2 | yes | |
| CNOT3 | CNOT3 | CCR4-Not Transcription Complex Subunit 3 | yes | |
| CNOT4 | CNOT4 | CCR4-Not Transcription Complex Subunit 4 | yes | |
| CNOT6 | CNOT6 | CCR4-Not Transcription Complex Subunit 6 | yes | |
| CNOT6L | CNOT6L | CCR4-Not Transcription Complex Subunit 6L | yes | |
| CNOT7 | CNOT7 | CCR4-Not Transcription Complex Subunit 7 | yes | |
| CNOT8 | CNOT8 | CCR4-Not Transcription Complex Subunit 8 | yes | |
| CNOT9 | CNOT9 | CCR4-Not Transcription Complex Subunit 9 | ||
| CORO1B | CORO1B | Coronin-1B | ||
| CPB2 | Carboxypeptidase B2 | Carboxypeptidase B2 | ||
| CPEB1 | CPEB | Cytoplasmic Polyadenylation Element Binding Protein 1 | ||
| CPEB4 | CPEB4 | Cytoplasmic Polyadenylation Element Binding Protein 4 | yes | |
| CPSF3 | CPSF3 | Cleavage and polyadenylation specificity factor subunit 3 | ||
| CPSF6 | CPSF6 | Cleavage and polyadenylation specificity factor subunit 6 | ||
| CPSF7 | CPSF7 | Cleavage and polyadenylation specificity factor subunit 7 | ||
| CPVL | CPVL | Carboxypeptidase, Vitellogenic Like | yes | |
| CRKL | CRKL | CRK Like Proto-Oncogene, Adaptor Protein | ||
| CROCC | CROCC | Ciliary Rootlet Coiled-Coil, Rootletin | ||
| CRYAB | CRYAB | Alpha-crystallin B chain | ||
| CSDE1 | CSDE1 | Cold shock domain-containing protein E1 | ||
| CSE1L | CSE1L/XPO2/Exportin-2 | Exportin-2 | ||
| CSNK2A1 | Casein Kinase 2 alpha | Casein Kinase 2 Alpha 1 | ||
| CSTB | Cystatin B | Cystatin B | ||
| CSTF1 | CSTF1 | Cleavage stimulation factor subunit 1 | ||
| CTNNA2 | CTNNA2 | Catenin alpha-2 | ||
| CTNND1 | CTNND1 | Catenin delta-1 | ||
| CTTNBP2NL | CTTNBP2NL | CTTNBP2 N-terminal-like protein | ||
| CWC22 | CWC22 | Pre-mRNA-splicing factor CWC22 homolog | ||
| DAZAP1 | DAZAP1 | DAZ-associated protein 1 | ||
| DAZAP2 | PRTB | DAZ Associated Protein 2 | ||
| DAZL | DAZL1 | Deleted In Azoospermia Like | ||
| DCD | DCD | Dermcidin | ||
| DCP1A | DCP1a | Decapping mRNA 1a | yes | |
| DCP1B | DCP1b | Decapping mRNA 1b | yes | |
| DCP2 | DCP2 | Decapping mRNA 2 | ||
| DCTN1 | DCTN1 | Dynactin subunit 1 | ||
| DDX1 | DEAD box protein 1 | DEAD-Box Helicase 1 | ||
| DDX19A | DDX19A | ATP-dependent RNA helicase DDX19A | ||
| DDX21 | DDX21 | Nucleolar RNA helicase 2 | yes | |
| DDX3 | DEAD box protein 3 | DEAD-Box Helicase 3 | ||
| DDX3X | DDX3X | DEAD-Box Helicase 3, X-Linked | ||
| DDX3Y | DDX3Y | DEAD-Box Helicase 3, Y-Linked | ||
| DDX47 | DDX47 | Probable ATP-dependent RNA helicase DDX47 | ||
| DDX50 | DDX50 | ATP-dependent RNA helicase DDX50 | yes | |
| DDX58 | RIG-I | DExD/H-Box Helicase 58 | ||
| DDX6 | DEAD box protein 6 | DEAD-Box Helicase 6 | yes | |
| DERA | DERA | Deoxyribose-Phosphate Aldolase | ||
| DHX30 | DHX30 | Putative ATP-dependent RNA helicase DHX30 | yes | |
| DHX33 | DHX33 | DEAH-Box Helicase 33 | ||
| DHX36 | RHAU | DEAH-Box Helicase 36 | ||
| DHX57 | DHX57 | DExH-Box Helicase 57 | ||
| DHX58 | LGP2 | DExH-Box Helicase 58 | ||
| DIS3L2 | DIS3L2/FAM3A | DIS3 Like 3'-5' Exoribonuclease 2 | ||
| DISC1 | Disrupted in Schizophrenia 1 | Disrupted In Schizophrenia 1 | ||
| DKC1 | DKC1 | dyskerin; H/ACA ribonucleoprotein complex subunit 4 | ||
| DNAI1 | Axonemal Dynein Intermediate Chain 1 | Dynein Axonemal Intermediate Chain 1 | ||
| DNAJA1 | DNAJA1 | DnaJ homolog subfamily A member 1 | ||
| DNAJC8 | DNAJC8 | DnaJ homolog subfamily C member 8 | ||
| DPYSL2 | DPYSL2 | Dihydropyrimidinase-related protein 2 | ||
| DPYSL3 | DPYSL3 | Dihydropyrimidinase-related protein 3 | ||
| DROSHA | DROSHA | Drosha Ribonuclease III | ||
| DSP | DSP | Desmoplakin | ||
| DST | DST | Dystonin | ||
| DSTN | DSTN | Destrin | ||
| DTX3L | DTX3L | E3 ubiquitin-protein ligase DTX3L | ||
| DUSP12 | DUSP12/YVH1 | Dual Specificity Phosphatase 12 | ||
| DYNC1H1 | Cytoplasmic Dynein Heavy Chain 1 | Dynein Cytoplasmic 1 Heavy Chain 1 | ||
| DYNLL1 | Cytoplasmic Dynein Light Polypeptide | Dynein Light Chain LC8-Type 1 | ||
| DYNLL2 | DYNLL2 | Dynein light chain 2, cytoplasmic | ||
| DYRK3 | DYRK3 | Dual Specificity Tyrosine Phosphorylation Regulated Kinase 3 | ||
| DZIP1 | DZIP1 | DAZ Interacting Zinc Finger Protein 1 | ||
| DZIP3 | DZIP3 | DAZ Interacting Zinc Finger Protein 3 | ||
| EDC3 | EDC3 | Enhancer of mRNA Decapping 3 | yes | |
| EDC4 | EDC4 | Enhancer of mRNA-Decapping protein 4 | yes | |
| EIF1 | EIF1 | Eukaryotic Translation Initiation Factor 1 | ||
| EIF2A | EIF2A | Eukaryotic Translation Initiation Factor 2A | ||
| EIF2AK2 | Protein Kinase R/PKR | Eukaryotic Translation Initiation Factor 2 Alpha Kinase 2 | ||
| EIF2B1-5 | EIF2B | Eukaryotic Translation Initiation Factor 2B | ||
| EIF2S1 | EIF2A subunit 1 | Eukaryotic Translation Initiation Factor 2 Subunit Alpha | ||
| EIF2S2 | EIF2A subunit 2 | Eukaryotic Translation Initiation Factor 2 Subunit Beta | ||
| EIF3A | EIF3A | Eukaryotic Translation Initiation Factor 3 Subunit A | ||
| EIF3B | EIF3B | Eukaryotic Translation Initiation Factor 3 Subunit B | ||
| EIF3C | EIF3C | Eukaryotic Translation Initiation Factor 3 Subunit C | ||
| EIF3D | EIF3D | Eukaryotic translation initiation factor 3 subunit D | ||
| EIF3E | EIF3E | Eukaryotic translation initiation factor 3 subunit E | ||
| EIF3F | EIF3F | Eukaryotic translation initiation factor 3 subunit F | ||
| EIF3G | EIF3G | Eukaryotic translation initiation factor 3 subunit G | ||
| EIF3H | EIF3H | Eukaryotic translation initiation factor 3 subunit H | ||
| EIF3I | EIF3I | Eukaryotic translation initiation factor 3 subunit I | ||
| EIF3J | EIF3J | Eukaryotic translation initiation factor 3 subunit J | ||
| EIF3K | EIF3K | Eukaryotic translation initiation factor 3 subunit K | ||
| EIF3L | EIF3L | Eukaryotic translation initiation factor 3 subunit L | ||
| EIF3M | EIF3M | Eukaryotic translation initiation factor 3 subunit M | ||
| EIF4A1 | EIF4A1 | Eukaryotic Translation Initiation Factor 4A1 | ||
| EIF4A2 | EIF4A2 | Eukaryotic Translation Initiation Factor 4A2 | ||
| EIF4A3 | EIF4A3 | Eukaryotic Translation Initiation Factor 4A3 | ||
| EIF4B | EIF4B | Eukaryotic translation Initiation factor 4B | ||
| EIF4E | EIF4E | Eukaryotic Translation Initiation Factor 4E | yes | |
| EIF4E2 | EIF4E2 | Eukaryotic Translation Initiation Factor 4E Family Member 2 | yes | |
| EIF4E3 | EIF4E3 | Eukaryotic Translation Initiation Factor 4E Family Member 3 | ||
| EIF4ENIF1 | EIF4ENIF1 | Eukaryotic Translation Initiation Factor 4E Nuclear Import Factor 1 | yes | |
| EIF4G1 | EIF4G1 | Eukaryotic Translation Initiation Factor 4G1 | ||
| EIF4G2 | EIF4G2 | Eukaryotic Translation Initiation Factor 4G2 | ||
| EIF4G3 | EIF4G3 | Eukaryotic Translation Initiation Factor 4G3 | ||
| EIF4H | EIF4H | Eukaryotic translation Initiation factor 4H | ||
| EIF5A | EIF5A | Eukaryotic Translation Initiation Factor 5A | ||
| ELAVL1 | HuR | ELAV Like RNA Binding Protein 1 | yes | |
| ELAVL2 | ELAVL2 | ELAV-like protein 2 | yes | |
| ELAVL3 | ELAVL3/HuC | ELAV Like RNA Binding Protein 3 | ||
| ELAVL4 | HuD | ELAV Like RNA Binding Protein 4 | ||
| ENDOV | EndoV | Endonuclease V | ||
| ENTPD1 | ENTPD1 | Ectonucleoside Triphosphate Diphosphohydrolase 1 | ||
| EPPK1 | EPPK1 | Epiplakin | ||
| ETF1 | ETF1 | Eukaryotic peptide chain release factor subunit 1 | ||
| EWSR1 | EWSR1 | EWS RNA Binding Protein 1 | ||
| FABP5 | FABP5 | Fatty Acid Binding Protein 5 | ||
| FAM120A | FAM120A/OSSA | Constitutive coactivator of PPAR-gamma-like protein 1 | yes | |
| FAM120C | FAM120C | Family With Sequence Similarity 120C | ||
| FAM168B | FAM168B/MANI | Family With Sequence Similarity 168 Member B | ||
| FAM98A | FAM98A | Family With Sequence Similarity 98 Member A | ||
| FASTK | FAST | Fas Activated Serine/Threonine Kinase | yes | |
| FBL | FBL | rRNA 2-O-methyltransferase fibrillarin | ||
| FBRSL1 | Fibrosin Like 1 | Fibrosin Like 1 | ||
| FHL1 | FHL1 | Four and a half LIM domains protein 1 | ||
| FLNB | FLNB | Filamin-B | ||
| FMR1 | FMRP | Fragile X Mental Retardation 1 | ||
| FNDC3B | FNDC3B | Fibronectin type III domain-containing protein 3B | ||
| FSCN1 | FSCN1 | Fascin | ||
| FTSJ3 | FTSJ3 | pre-rRNA processing protein FTSJ3 | ||
| FUBP1 | FUBP1 | Far Upstream Element Binding Protein 1 | ||
| FUBP3 | FUBP3 | Far upstream element-binding protein 3 | ||
| FUS | FUS | FUS RNA Binding Protein | ||
| FXR1 | FXR1 | FMR1 Autosomal Homolog 1 | ||
| FXR2 | FXR2 | FMR1 Autosomal Homolog 2 | ||
| G3BP1 | G3BP1 | G3BP Stress Granule Assembly Factor 1 | ||
| G3BP2 | G3BP2 | G3BP Stress Granule Assembly Factor 2 | ||
| GABARAPL2 | GABARAPL2/GEF2/ATG8 | GABA Type A Receptor Associated Protein Like 2 | ||
| GAR1 | GAR1 | H/ACA Ribonucleoprotein Complex Subunit 1 | ||
| GCA | Grancalcin | Grancalcin | ||
| GEMIN5 | Gemin-5 | Gem Nuclear Organelle Associated Protein 5 | ||
| GFPT1 | GFPT1 | Glutamine—fructose-6-phosphate aminotransferase 1 | ||
| GIGYF1 | GIGYF1/PERQ1 | GRB10 Interacting GYF Protein 1 | ||
| GIGYF2 | GIGYF2/TNRC15/PARK11/PERQ2 | GRB10 Interacting GYF Protein 2 | yes | |
| GLE1 | GLE1 | GLE1, RNA Export Mediator | ||
| GLO1 | Glyoxalase | Glyoxalase | ||
| GLRX3 | GLRX3/Glutaredoxin 3/TNLX2 | Glutaredoxin 3 | ||
| GNB2 | GNB2 | Guanine nucleotide-binding protein G/G/G subunit beta-2 | ||
| GOLGA2 | Golgin A2 | Golgin A2 | ||
| GRB2 | GRB2/ASH | Growth Factor Receptor Bound Protein 2 | ||
| GRB7 | GRB7 | Growth Factor Receptor Bound Protein 7 | ||
| GRSF1 | GRSF1 | G-Rich RNA Sequence Binding Factor 1 | ||
| GSPT1 | eRF3 | G1 To S Phase Transition 1 | ||
| H1F0 | H1F0 | Histone H1.0 | ||
| H1FX | H1FX | Histone H1x | ||
| H2AFV | H2AFV | Histone H2A.V | ||
| HABP4 | Ki-1/57 | Hyaluronan Binding Protein 4 | ||
| HDAC6 | HDAC6 | Histone Deacetylase 6 | ||
| HDLBP | HDL-Binding Protein/VGL/Vigilin | High Density Lipoprotein Binding Protein | ||
| HELZ | HELZ | Probable helicase with zinc finger domain | yes | |
| HELZ2 | HELZ2 | Helicase with zinc finger domain 2 | ||
| HMGA1 | HMGA1 | High mobility group protein HMG-I/HMG-Y | ||
| HMGB3 | HMGB3 | High mobility group protein B3 | ||
| HMGN1 | HMGN1 | Non-histone chromosomal protein HMG-14 | ||
| HNRNPA1 | HnRNPA1 | Heterogeneous Nuclear Ribonucleoprotein A1 | ||
| HNRNPA2B1 | HnRNPA2/B1 | Heterogeneous Nuclear Ribonucleoprotein A2/B1 | ||
| HNRNPA3 | HNRNPA3 | Heterogeneous nuclear ribonucleoprotein A3 | ||
| HNRNPAB | HNRNPAB | Heterogeneous nuclear ribonucleoprotein A/B | ||
| HNRNPD | HNRNPD | Heterogeneous nuclear ribonucleoprotein D | ||
| HNRNPDL | HNRNPDL | Heterogeneous nuclear ribonucleoprotein D-like | ||
| HNRNPF | HNRNPF | Heterogeneous nuclear ribonucleoprotein F | ||
| HNRNPH1 | HNRNPH1 | Heterogeneous nuclear ribonucleoprotein H1 | ||
| HNRNPH2 | HNRNPH2 | Heterogeneous nuclear ribonucleoprotein H2 | ||
| HNRNPH3 | HNRNPH3 | Heterogeneous nuclear ribonucleoprotein H3 | ||
| HNRNPK | HNRNPK | Heterogeneous Nuclear Ribonucleoprotein K | ||
| HNRNPUL1 | HNRNPUL1 | Heterogeneous nuclear ribonucleoprotein U-like protein 2 | ||
| HSBP1 | HSBP1 | Heat Shock Factor Binding Protein 1 | ||
| HSP90AA1 | HSP90 | Heat shock protein HSP 90-alpha | ||
| HSPA4 | HSP70 RY | Heat shock 70 kDa protein 4 | ||
| HSPA9 | HSP70 9B | Stress-70 protein, mitochondrial | ||
| HSPB1 | HSP27 | Heat Shock Protein Family B Member 1 | yes | |
| HSPB8 | HSPB8 | Heat Shock Protein Family B Member 8 | ||
| HSPBP1 | HSPBP1 | HSPA Binding Protein 1 | ||
| HSPD1 | HSPD1 | 60 kDa heat shock protein, mitochondrial | ||
| HTT | Huntingtin | Huntingtin | ||
| IBTK | IBTK | Inhibitor Of Bruton Tyrosine Kinase | ||
| IFIH1 | MDA5 | Interferon Induced With Helicase C Domain 1 | ||
| IGF2BP1 | IGF2BP1 | Insulin-like Growth Factor 2 mRNA-binding protein 1 | yes | |
| IGF2BP2 | IGF2BP2 | Insulin-like Growth Factor 2 mRNA-binding protein 2 | yes | |
| IGF2BP3 | IGF2BP3 | Insulin-like Growth Factor 2 mRNA Binding Protein 3 | yes | |
| IK | IK | Protein Red | ||
| ILF3 | NF90 | Interleukin Enhancer Binding Factor 3 | yes | |
| IPO7 | IPO7 | Importin-7 | ||
| IPPK | IP5K | Inositol-Pentakisphosphate 2-Kinase | ||
| ITGB1 | ITGB1 | Integrin beta-1 | ||
| JMJD6 | JMJD6 | Arginine Demethylase and Lysine Hydroxylase | ||
| KANK2 | KANK2 | KN motif and ankyrin repeat domain-containing protein 2 | ||
| KEAP1 | KEAP1/KLHL19 | Kelch Like ECH Associated Protein 1 | ||
| KHDRBS1 | Sam68 | KH RNA Binding Domain Containing, Signal Transduction Associated 1 | ||
| KHDRBS3 | KHDRBS3 | KH domain-containing, RNA-binding, signal transduction-associated protein 3 | ||
| KHSRP | KSRP/FBP2 | KH-Type Splicing Regulatory Protein | ||
| KIAA0232 | KIAA0232 | KIAA0232 | yes | |
| KIAA1524 | CIP2A | Protein CIP2A | ||
| KIF1B | KIF1B | Kinesin Family Member 1B | ||
| KIF13B | KIF13B/GAKIN | Kinesin Family Member 13B | ||
| KIF23 | KIF23 | Kinesin-like protein KIF23 | yes | |
| KIF2A | Kinesin Heavy Chain Member 2 | Kinesin Family Member 2A | ||
| KLC1 | Kinesin Light Chain 1 | Kinesin Light Chain 1 | ||
| KPNA1 | Importin-ɑ5 | Karyopherin Subunit Alpha 1 | ||
| KPNA2 | Importin-ɑ1 | Karyopherin Subunit Alpha 2 | ||
| KPNA3 | Importin-ɑ4 | Karyopherin Subunit Alpha 3 | ||
| KPNA6 | Importin-ɑ7 | Importin subunit alpha | ||
| KPNB1 | Importin-β1 | Karyopherin Subunit Beta 1 | ||
| L1RE1 | LINE1 ORF1p | LINE1 ORF1 protein | ||
| LANCL1 | LanC Like 1 | LanC Like 1 | ||
| LARP1 | LARP1 | La-related protein 1 | ||
| LARP1B | LARP1B | La-related protein 1b | ||
| LARP4 | La-Related protein 4 | La Ribonucleoprotein Domain Family Member 4 | ||
| LARP4B | LARP4B | La Ribonucleoprotein Domain Family Member 4B | ||
| LASP1 | LIM And SH3 Protein 1/MLN50 | LIM And SH3 Protein 1 | ||
| LBR | LBR | Lamin-B receptor | ||
| LEMD3 | LEMD3 | Inner nuclear membrane protein Man1 | ||
| LIG3 | DNA Ligase 3 | DNA Ligase 3 | ||
| LIN28A | LIN28A | Lin-28 Homolog A | ||
| LIN28B | LIN28B | Lin-28 Homolog B | ||
| LMNA | LMNA | Prelamin-A/C | ||
| LPP | LPP | Lipoma-preferred partner | ||
| LSM1 | LSM1 | LSM1 Homolog, mRNA Degradation Associated | yes | |
| LSM12 | LSM12 | LSM12 Homolog | ||
| LSM14A | RAP55 | LSM14A, mRNA Processing Body Assembly Factor | yes | |
| LSM14B | LSM14B | Protein LSM14 homolog B | yes | |
| LSM3 | LSM3 | U6 snRNA-associated Sm-like protein LSm3 | yes | |
| LUC7L | LUC7L | Putative RNA-binding protein Luc7-like 1 | ||
| LUZP1 | LUZP1 | Leucine zipper protein 1 | ||
| MACF1 | MACF1 | Microtubule-actin cross-linking factor 1, isoforms 1/2/3/5 | ||
| MAEL | MAEL | Maelstrom Spermatogenic Transposon Silencer | ||
| MAGEA4 | MAGEA4 | Melanoma-associated antigen 4 | ||
| MAGED1 | MAGED1 | Melanoma-associated antigen D1 | ||
| MAGED2 | MAGED2 | Melanoma-associated antigen D2 | ||
| MAGOHB | MAGOHB | Protein mago nashi homolog 2 | ||
| MAP1LC3A | LC3-I | Microtubule Associated Protein 1 Light Chain 3 Alpha | ||
| MAP4 | MAP4 | Microtubule-associated protein 4 | ||
| MAPK1IP1L | MAPK1IP1L | Mitogen-Activated Protein Kinase 1 Interacting Protein 1 Like | ||
| MAP4K4 | MAP4K4 | Mitogen-activated protein kinase kinase kinase kinase 4 | ||
| MAPK8 | JNK1 | Mitogen-Activated Protein Kinase 8 | ||
| MAPRE1 | MAPRE1 | Microtubule-associated protein RP/EB family member 1 | ||
| MAPRE2 | MAPRE2 | Microtubule Associated Protein RP/EB Family Member 2 | ||
| MARF1 | MARF1 | Meiosis Regulator And mRNA Stability Factor 1 | yes | |
| MARS | MARS | Methionine—tRNA ligase, cytoplasmic | ||
| MBNL1 | MBNL1 | Muscleblind Like Splicing Regulator 1 | ||
| MBNL2 | MBNL2 | Muscleblind Like Splicing Regulator 2 | ||
| MCM4 | MCM4 | DNA replication licensing factor MCM4 | ||
| MCM5 | MCM5 | DNA replication licensing factor MCM5 | ||
| MCM7 | MCM7 | DNA replication licensing factor MCM7 | yes | |
| METAP1 | METAP1 | Methionine aminopeptidase | ||
| METAP2 | METAP2 | Methionyl Aminopeptidase 2 | ||
| MCRIP1 | FAM195B/GRAN2 | Granulin-2 | ||
| MCRIP2 | FAM195A/GRAN1 | Granulin-1 | ||
| MEX3A | MEX3A | RNA-binding protein MEX3A | yes | |
| MEX3B | MEX3B | Mex-3 RNA Binding Family Member B | ||
| MEX3C | MEX3C | Mex-3 RNA Binding Family Member C | ||
| MEX3D | MEX3D | Mex-3 RNA Binding Family Member D | ||
| MFAP1 | MFAP1 | Microfibrillar-associated protein 1 | ||
| MKI67 | MKI67 | Antigen KI-67 | ||
| MKRN2 | MKRN2 | Makorin Ring Finger Protein 2 | ||
| MOV10 | MOV-10 | Mov10 RISC Complex RNA Helicase | yes | |
| MSH6 | MSH6 | DNA mismatch repair protein Msh6 | ||
| MSI1 | Musashi-1 | Musashi RNA Binding Protein 1 | yes | |
| MSI2 | MSI2 | RNA-binding protein Musashi homolog 2 | ||
| MTHFD1 | MTHFD1 | C-1-tetrahydrofolate synthase, cytoplasmic | ||
| MTHFSD | MTHFSD | Methenyltetrahydrofolate Synthetase Domain Containing | ||
| MTOR | MTOR | Mechanistic Target Of Rapamycin | ||
| MYO6 | MYO6 | Unconventional myosin-VI | ||
| NCOA3 | SRC-3 | Nuclear Receptor Coactivator 3 | ||
| NDEL1 | NUDEL/MITAP1/EOPA | NudE Neurodevelopment Protein 1 Like 1 | ||
| NELFE | NELF-E/RD | Negative Elongation Factor Complex Member E | ||
| NEXN | NEXN | Nexilin | ||
| NXF1 | NXF1/MEX67/TAP | Nuclear RNA Export Factor 1 | ||
| NKRF | NRF | NFK-B Repressing Factor | ||
| NOLC1 | Nucleolar And Coiled-Body Phosphoprotein 1/NOPP140 | Nucleolar And Coiled-Body Phosphoprotein 1 | ||
| NONO | NonO | Non-POU Domain Containing Octamer Binding | ||
| NOP58 | NOP58 | Nucleolar protein 58 | yes | |
| NOSIP | NOSIP | Nitric oxide synthase-interacting protein | ||
| NOVA2 | NOVA2 | NOVA Alternative Splicing Regulator 2 | ||
| NRG2 | Neuregulin-2 | Neuregulin-2 | ||
| NSUN2 | NSUN2 | tRNA -C)-methyltransferase | ||
| NTMT1 | NTMT1 | N-terminal Xaa-Pro-Lys N-methyltransferase 1 | ||
| NUDC | NUDC | Nuclear migration protein nudC | ||
| NUFIP1 | NUFIP | NUFIP1, FMR1 Interacting Protein 1 | ||
| NUFIP2 | NUFIP2 | Nuclear fragile X mental retardation-interacting protein 2 | ||
| NUPL2 | NUPL2 | Nucleoporin Like 2 | ||
| NUP153 | NUP153 | Nucleoporin 153 | ||
| NUP205 | NUP205 | Nuclear pore complex protein Nup205 | ||
| NUP210 | NUP210/GP210 | Nucleoporin 210 | ||
| NUP214 | NUP214 | Nucleoporin 214 | ||
| NUP50 | NUP50 | Nucleoporin 50 | ||
| NUP58 | NUP58/NUPL1 | Nucleoporin 58 | ||
| NUP85 | NUP85 | Nucleoporin 85 | ||
| NUP88 | NUP88 | Nucleoporin 88 | ||
| NUP98 | NUP98/NUP96 | Nuclear pore complex protein Nup98-Nup96 | ||
| OASL | OASL/OASL1 | 2'-5'-Oligoadenylate Synthetase Like | ||
| OAS1 | OAS | 2′–5′ oligoadenylate synthetase | ||
| OAS2 | OAS2 | 2'-5'-Oligoadenylate Synthetase 2 | ||
| OGFOD1 | TPA1 | 2-Oxoglutarate And Iron Dependent Oxygenase Domain Containing 1 | ||
| OGG1 | OGG1 | 8-Oxoguanine DNA Glycosylase | ||
| OSBPL9 | Oxysterol Binding Protein Like 9 | Oxysterol Binding Protein Like 9 | ||
| OTUD4 | OTUD4/HIN1 | OTU Deubiquitinase 4 | ||
| P4HB | Prolyl 4-Hydroxylase Subunit Beta | Prolyl 4-Hydroxylase Subunit Beta | ||
| PABPC1 | PABP1 | Poly Binding Protein Cytoplasmic 1 | ||
| PABPC4 | PABPC4 | Polyadenylate-binding protein 4 | ||
| PAK4 | PAK4 | Serine/threonine-protein kinase PAK 4 | ||
| PALLD | Palladin | Palladin | ||
| PARG | PARG/PARG99/PARG102 | Poly Glycohydrolase | ||
| PARK7 | PARK7/DJ-1 | Parkinsonism Associated Deglycase | yes | |
| PARN | PARN/DAN | Poly-Specific Ribonuclease | ||
| PARP12 | PARP-12/ARTD12 | Poly Polymerase Family Member 12 | ||
| PARP14 | PARP-14 | Poly Polymerase Family Member 14 | ||
| PARP15 | PARP-15 | Poly Polymerase Family Member 15 | ||
| PATL1 | PATL1 | PAT1 Homolog 1, Processing Body mRNA Decay Factor | yes | |
| PAWR | PAWR | PRKC apoptosis WT1 regulator protein | ||
| PCBP1 | PCBP1/HNRNPE1 | Poly Binding Protein 1 | ||
| PCBP2 | PCBP2/HNRNPE2 | Poly Binding Protein 2 | ||
| PCNA | PCNA | Proliferating cell nuclear antigen | ||
| PDAP1 | PDAP1 | PDGFA Associated Protein 1 | ||
| PDCD4 | PDCD4 | Programmed Cell Death 4 | ||
| PDCD6IP | PDCD6IP | Programmed cell death 6-interacting protein | ||
| PDIA3 | PDIA3 | Protein Disulfide Isomerase Family A Member 3 | ||
| PDLIM1 | PDLIM1 | PDZ and LIM domain protein 1 | ||
| PDLIM4 | PDLIM4 | PDZ and LIM domain protein 4 | ||
| PDLIM5 | PDLIM5 | PDZ and LIM domain protein 5 | ||
| PDS5B | PDS5B | Sister chromatid cohesion protein PDS5 homolog B | ||
| PEF1 | PEF1 | Penta-EF-Hand Domain Containing 1 | ||
| PEG10 | PEG10 | Paternally Expressed 10 | ||
| PELO | PELO | Protein pelota homolog | ||
| PEPD | Peptidase D | Peptidase D | ||
| PEX11B | PEX11B | Peroxisomal Biogenesis Factor 11 Beta | ||
| PFDN4 | PFDN4 | Prefoldin subunit 4 | ||
| PFN1 | Profilin 1 | Profilin 1 | ||
| PFN2 | Profilin 2 | Profilin 2 | ||
| PGAM5 | PGAM5 | Serine/threonine-protein phosphatase PGAM5, mitochondrial | ||
| PGP | PGP/G3PP | Phosphoglycolate Phosphatase | ||
| PHB2 | Prohibitin 2 | Prohibitin 2 | ||
| PHLDB2 | PHLDB2 | Pleckstrin homology-like domain family B member 2 | ||
| PKP1 | Plakophilin 1 | Plakophilin 1 | ||
| PKP2 | Plakophilin 2 | Plakophilin 2 | ||
| PKP3 | Plakophilin 3 | Plakophilin 3 | ||
| PNPT1 | PNPase I | Polyribonucleotide Nucleotidyltransferase 1 | ||
| POLR2B | POLR2B | DNA-directed RNA polymerase | ||
| POM121 | POM121 | POM121 Transmembrane Nucleoporin | ||
| POP7 | RPP20 | POP7 Homolog, Ribonuclease P/MRP Subunit | ||
| PPME1 | PPME1 | Protein phosphatase methylesterase 1 | ||
| PPP1R8 | PPP1R8 | Protein Phosphatase 1 Regulatory Subunit 8 | ||
| PPP1R10 | PPP1R10 | Serine/threonine-protein phosphatase 1 regulatory subunit 10 | ||
| PPP1R18 | PPP1R18 | Phostensin | ||
| PPP2R1A | PPP2R1A | Serine/threonine-protein phosphatase 2A 65 kDa regulatory subunit A alpha isoform | ||
| PPP2R1B | PPP2R1B | Serine/threonine-protein phosphatase 2A 65 kDa regulatory subunit A beta isoform | ||
| PQBP1 | PQBP-1 | Polyglutamine Binding Protein 1 | ||
| PRDX1 | PRDX1 | Peroxiredoxin-1 | ||
| PRDX6 | PRDX6 | Peroxiredoxin-6 | ||
| PRKAA2 | AMPK-a2 | Protein Kinase AMP-Activated Catalytic Subunit Alpha 2 | ||
| PRKCA | PKC-ɑ | Protein Kinase C Alpha | ||
| PRKRA | PACT | Protein Activator Of Interferon Induced Protein Kinase EIF2AK2 | ||
| PRMT1 | PRMT1 | Protein arginine N-methyltransferase 1 | ||
| PRMT5 | PRMT5 | Protein arginine N-methyltransferase 5 | ||
| PRRC2A | PRRC2A | Proline Rich Coiled-Coil 2A | ||
| PRRC2B | PRRC2B | Proline Rich Coiled-Coil 2B | ||
| PRRC2C | PRRC2C | Proline Rich Coiled-Coil 2C | ||
| PSMD2 | PSMD2 | 26S proteasome non-ATPase regulatory subunit 2 | ||
| PSPC1 | PSP1 | Paraspeckle Component 1 | ||
| PTBP1 | PTBP1 | Polypyrimidine tract-binding protein 1 | ||
| PTBP3 | PTBP3 | Polypyrimidine tract-binding protein 3 | ||
| PTGES3 | PTGES3 | Prostaglandin E synthase 3 | ||
| PTK2 | FAK | Protein Tyrosine Kinase 2 | ||
| PUM1 | Pumilio-1 | Pumilio homolog 1 | yes | |
| PUM2 | Pumilio-2 | Pumilio RNA Binding Family Member 2 | ||
| PURA | PURA | Transcriptional activator protein Pur-alpha | ||
| PURB | PURB | Transcriptional activator protein Pur-beta | ||
| PWP1 | PWP1 | PWP1 Homolog, Endonuclein | ||
| PXDNL | PMR1 | Peroxidasin Like | ||
| PYCR1 | PYCR1 | Pyrroline-5-carboxylate reductase | ||
| QKI | QKI/HQK | QKI, KH Domain Containing RNA Binding | ||
| R3HDM1 | R3HDM1 | R3H Domain Containing 1 | ||
| R3HDM2 | R3HDM2 | R3H Domain Containing 2 | ||
| RAB1A | RAB1A | Ras-related protein Rab-1A | ||
| RACGAP1 | RACGAP1 | Rac GTPase-activating protein 1 | ||
| RACK1 | RACK1 | Receptor For Activated C Kinase 1 | ||
| RAD21 | RAD21 | Double-strand-break repair protein rad21 homolog | ||
| RAE1 | RAE1 | Ribonucleic Acid Export 1 | ||
| RAN | RAN | RAN, Member RAS Oncogene Family | ||
| RANBP1 | RANBP1 | Ran-specific GTPase-activating protein | ||
| RANBP2 | RANBP2/NUP358 | RAN Binding Protein 2 | ||
| RBBP4 | RBBP4 | Histone-binding protein RBBP4 | ||
| RBFOX1 | RBFOX1 | RNA binding protein fox-1 homolog | yes | |
| RBFOX2 | RBFOX2 | RNA binding protein fox-1 homolog 2 | ||
| RBFOX3 | RBFOX3 | RNA binding protein fox-1 homolog 3 | ||
| RBM12B | RBM12B | RNA-binding protein 12B | ||
| RBM15 | RBM15 | RNA-binding protein 15 | ||
| RBM17 | RBM17 | RNA-binding protein 17 | ||
| RBM25 | RBM25 | RNA-binding protein 25 | ||
| RBM26 | RBM26 | RNA-binding protein 26 | ||
| RBM3 | RBM3 | RNA-binding protein 3 | ||
| RBM38 | RBM38 | RNA-binding protein 38 | ||
| RBM4 | RBM4 | RNA Binding Motif Protein 4 | ||
| RBM4B | RBM4B | RNA Binding Motif Protein 4B | ||
| RBM42 | RBM42 | RNA Binding Motif Protein 42 | ||
| RBM45 | RBM45 | RNA Binding Motif Protein 45 | ||
| RBM47 | RBM47 | RNA Binding Motif Protein 47 | ||
| RBMS1 | RBMS1 | RNA-binding motif, single-stranded-interacting protein 1 | ||
| RBMS2 | RBMS2 | RNA-binding motif, single-stranded-interacting protein 2 | ||
| RBMX | RBMX | RNA Binding Motif Protein, X-Linked | ||
| RBPMS | RBPMS | RNA-binding protein with multiple splicing | ||
| RC3H1 | Roquin-1 | Ring Finger And CCCH-Type Domains 1 | ||
| RC3H2 | MNAB | Ring Finger And CCCH-Type Domains 2 | ||
| RCC1 | RCC1 | Regulator of chromosome condensation | ||
| RCC2 | RCC2 | Protein RCC2 | ||
| RECQL | RECQL1 | RecQ Like Helicase | ||
| RFC3 | RFC3 | Replication factor C subunit 3 | ||
| RFC4 | RFC4 | Replication factor C subunit 4 | ||
| RGPD3 | RGPD3 | RanBP2-like and GRIP domain-containing protein 3 | ||
| RHOA | RhoA | Ras Homolog Family Member A | ||
| RNASEL | RNAse L | Ribonuclease L | ||
| RNF214 | RNF214 | RING finger protein 214 | ||
| RNF219 | RNF219 | RING finger protein 219 | yes | |
| RNF25 | RNF25 | Ring Finger Protein 25 | ||
| RNH1 | RNH1 | Ribonuclease inhibitor | ||
| ROCK1 | ROCK1 | Rho Associated Coiled-Coil Containing Protein Kinase 1 | ||
| RPS19 | Ribosomal Protein S19 | Ribosomal Protein S19 | ||
| RPS3 | 40S Ribosomal Protein S3 | 40S Ribosomal Protein S3 | yes | |
| RPS6 | Ribosomal Protein S6 | Ribosomal Protein S6 | ||
| RPS11 | Ribosomal Protein S11 | Ribosomal Protein S11 | ||
| RPS24 | Ribosomal Protein S24 | Ribosomal Protein S24 | ||
| RPS6KA3 | RSK2 | Ribosomal Protein S6 Kinase A3 | ||
| RPS6KB1 | S6K1 | Ribosomal Protein S6 Kinase B1 | ||
| RPS6KB2 | S6K2 | Ribosomal Protein S6 Kinase B2 | ||
| RPTOR | RAPTOR | Regulatory Associated Protein of mTOR Complex 1 | ||
| RSL1D1 | RSL1D1 | Ribosomal L1 domain-containing protein 1 | ||
| RTCB | RTCB | tRNA-splicing ligase RtcB homolog, formerly C22orf28 | ||
| RTRAF | RTRAF | RNA Transcription, Translation And Transport Factor | ||
| S100A7A | S100A7A | Protein S100-A7A | ||
| S100A9 | S100A9 | Protein S100-A9 | yes | |
| SAFB2 | SAFB2 | Scaffold attachment factor B2 | yes | |
| SAMD4A | SMAUG1 | Sterile Alpha Motif Domain Containing 4A | ||
| SAMD4B | SMAUG2 | Sterile Alpha Motif Domain Containing 4B | ||
| SCAPER | SCAPER | S-Phase Cyclin A Associated Protein In The ER | ||
| SEC24C | SEC24C | Protein transport protein Sec24C | ||
| SECISBP2 | SECIS Binding Protein 2 | SECIS Binding Protein 2 | ||
| SERBP1 | PAI-RBP1/SERBP1 | SERPINE1 mRNA Binding Protein 1 | ||
| SERPINE1 | PAI-1/Serpin E1 | Serpine Family E Member 1 | ||
| SF1 | SF1 | Splicing Factor 1 | ||
| SFN | SFN | 14-3-3 protein sigma | ||
| SFPQ | PSF | Splicing Factor Proline And Glutamine Rich | ||
| SFRS3 | SFRS3 | Serine/arginine-rich splicing factor 3 | ||
| SIPA1L1 | SIPA1L1 | Signal-induced proliferation-associated 1-like protein 1 | ||
| SIRT6 | Sirtuin 6 | Sirtuin 6 | ||
| SLBP | Stem-Loop Binding Protein | Stem-Loop Binding Protein | ||
| SMAP2 | SMAP2 | Small ArfGAP2 | ||
| SMARCA1 | SMARCA1/SNF2L1 | Probable global transcription activator SNF2L1 | ||
| SMC4 | SMC4 | Structural maintenance of chromosomes protein | ||
| SMG1 | SMG-1 | SMG1, Nonsense Mediated mRNA Decay Associated PI3K Related Kinase | ||
| SMG6 | SMG6 | SMG6, Nonsense Mediated mRNA Decay Factor | ||
| SMG7 | SMG7 | SMG7, Nonsense Mediated mRNA Decay Factor | yes | |
| SMN1 | Survival of Motor Neuron | Survival Of Motor Neuron 1, Telomeric | ||
| SMU1 | SMU1 | WD40 repeat-containing protein SMU1 | ||
| SMYD5 | SMYD5 | SMYD Family Member 5 | ||
| SND1 | Tudor-SN | Staphylococcal Nuclease And Tudor Domain Containing 1 | ||
| SNRPF | SNRPF | Small nuclear ribonucleoprotein F | ||
| SNTB2 | SNTB2 | Beta-2-syntrophin | ||
| SOGA3 | SOGA3 | SOGA Family Member 3 | ||
| SORBS1 | SORBS1 | Sorbin and SH3 domain-containing protein 1 | ||
| SORBS3 | Vinexin | Sorbin And SH3 Domain Containing 3 | ||
| SOX3 | SOX3 | SRY-Box 3 | ||
| SPAG5 | Astrin | Sperm Associated Antigen 5 | ||
| SPATS2 | SPATS2/SPATA10/SCR59 | Spermatogenesis Associated Serine Rich 2 | ||
| SPATS2L | SGNP | Spermatogenesis Associated Serine Rich 2 Like | ||
| SPECC1L | SPECC1L | Cytospin-A | ||
| SQSTM1 | SQSTM1/p62 | Sequestosome 1 | ||
| SRI | SRI | Sorcin | ||
| SRP68 | Signal Recognition Particle 68 | Signal Recognition Particle 68 | ||
| SRP9 | SRP9 | Signal Recognition Particle 9 | ||
| SRRT | SRRT | Serrate RNA effector molecule homolog | ||
| SRSF1 | ASF/SF2 | Serine And Arginine Rich Splicing Factor 1 | ||
| SRSF3 | SRp20 | Serine And Arginine Rich Splicing Factor 3 | ||
| SRSF4 | SRSF4 | Serine/arginine-rich splicing factor 4 | ||
| SRSF5 | SRSF5/SRP40 | Serine/arginine-rich splicing factor 5 | ||
| SRSF7 | 9G8 | Serine And Arginine Rich Splicing Factor 7 | ||
| SRSF9 | SRSF9/SRP30C | Serine/arginine-rich splicing factor 9 | ||
| SS18L1 | SS18L1/CREST | SS18L1, nBAF Chromatin Remodeling Complex Subunit | ||
| ST7 | ST7/FAM4A1/HELG/RAY1/TSG7 | Suppression Of Tumorigenicity 7 | yes | |
| STAT1 | STAT1 | Signal transducer and activator of transcription 1-alpha/beta | ||
| STAU1 | Staufen 1 | Staufen Double-Stranded RNA Binding Protein 1 | ||
| STAU2 | Staufen 2 | Staufen Double-Stranded RNA Binding Protein 2 | yes | |
| STIP1 | STIP1/HOP | Stress-induced-phosphoprotein 1 | ||
| STRAP | STRAP | Serine-threonine kinase receptor-associated protein | ||
| SUGP2 | SUGP2 | SURP and G-patch domain-containing protein 2 | ||
| SUGT1 | SUGT1 | SGT1 Homolog, MIS12 Kinetochore Complex Assembly Cochaperone | ||
| SUN1 | SUN1 | SUN domain-containing protein 1 | ||
| SYCP3 | SYCP3 | Synaptonemal complex protein 3 | ||
| SYK | SYK | Spleen Associated Tyrosine Kinase | ||
| SYNCRIP | SYNCRIP | Heterogeneous nuclear ribonucleoprotein Q | yes | |
| TAGLN3 | Transgelin 3 | Transgelin 3 | ||
| TAF15 | TAF15 | TATA-Box Binding Protein Associated Factor 15 | ||
| TARDBP | TDP-43 | TAR DNA Binding Protein | ||
| TBRG1 | TBRG1 | Transforming Growth Factor Beta Regulator 1 | ||
| TCEA1 | TCEA1 | Transcription elongation factor A protein 1 | ||
| TCP1 | TCP1 | T-complex protein 1 subunit alpha | ||
| TDRD3 | Tudor Domain Containing 3 | Tudor Domain Containing 3 | ||
| TDRD7 | Tudor Domain Containing 7 | Tudor Domain Containing 7 | ||
| TERT | TERT | Telomerase Reverse Transcriptase | ||
| THOC2 | THOC2 | THO Complex 2 | ||
| THRAP3 | THRAP3 | Thyroid Hormone Receptor Associated Protein 3 | ||
| TIA1 | TIA-1 | TIA1 Cytotoxic Granule Associated RNA Binding Protein | ||
| TIAL1 | TIAR | TIA1 Cytotoxic Granule Associated RNA Binding Protein Like 1 | ||
| TMEM131 | TMEM131 | Transmembrane Protein 131 | yes | |
| TMOD3 | TMOD3 | Tropomodulin-3 | ||
| TNKS | PARP-5a | Tankyrase | ||
| TNKS1BP1 | TNKS1BP1 | 182 kDa tankyrase-1-binding protein | yes | |
| TNPO1 | Transportin-1 | Transportin-1/Karyopherin Beta 2 | ||
| TNPO2 | Transportin-2 | Transportin-2 | ||
| TNRC6A | TNRC6A | Trinucleotide repeat-containing gene 6A protein | yes | |
| TNRC6B | TNRC6B | Trinucleotide repeat-containing gene 6B protein | yes | |
| TNRC6C | TNRC6C | Trinucleotide repeat-containing gene 6C protein | yes | |
| TOMM34 | TOMM34 | Mitochondrial import receptor subunit TOM34 | ||
| TOP3B | Topoisomerase III Beta | Topoisomerase III Beta | ||
| TPM1 | TPM1 | Tropomyosin alpha-1 chain | ||
| TPM2 | TPM2 | Tropomyosin beta chain | ||
| TPR | TPR | Translocated Promoter Region, Nuclear Basket Protein | ||
| TRA2B | TRA2B | Transformer 2 Beta Homolog | ||
| TRAF2 | TRAF2 | TNF Receptor Associated Factor 2 | ||
| TRDMT1 | DNMT2 | tRNA Aspartic Acid Methyltransferase 1 | ||
| TRIM21 | TRIM21 | E3 ubiquitin-protein ligase TRIM21 | ||
| TRIM25 | TRIM25 | E3 ubiquitin/ISG15 ligase TRIM25 | ||
| TRIM56 | TRIM56 | E3 ubiquitin-protein ligase TRIM56 | ||
| TRIM71 | TRIM71 | E3 ubiquitin-protein ligase TRIM71 | ||
| TRIP6 | TRIP6 | Thyroid receptor-interacting protein 6 | ||
| TROVE2 | RORNP | TROVE Domain Family Member 2 | ||
| TTC17 | TTC17 | Tetratricopeptide Repeat Domain 17 | yes | |
| TUBA1C | TUBA1C | Tubulin alpha-1C chain | ||
| TUBA3C | TUBA3C | Tubulin alpha-3C/D chain | ||
| TUBA4A | TUBA4A | Tubulin alpha-4A chain | ||
| TUBB3 | TUBB3 | Tubulin beta-3 chain | ||
| TUBB8 | TUBB8 | Tubulin beta-8 chain | ||
| TUFM | TUFM | Elongation factor Tu, mitochondrial | ||
| TXN | TXN | Thioredoxin | ||
| TXNDC17 | TXNDC17 | Thioredoxin Domain Containing 17 | ||
| U2AF1 | U2AF1 | Splicing factor U2AF 35 kDa subunit | ||
| UBA1 | UBA1 | Ubiquitin-like modifier-activating enzyme 1 | ||
| UBAP2 | UBAP2 | Ubiquitin-associated protein 2 | ||
| UBAP2L | UBAP2L | Ubiquitin-associated protein 2-like | ||
| UBB | Ubiquitin | Ubiquitin | ||
| UBL5 | Ubiquitin Like 5 | Ubiquitin Like 5 | ||
| UBQLN2 | Ubiquilin 2 | Ubiquilin 2 | ||
| ULK1 | ULK1 | Unc-51 Like Autophagy Activating Kinase 1 | ||
| ULK2 | ULK2 | Unc-51 Like Autophagy Activating Kinase 2 | ||
| UPF1 | UPF1 | UPF1, RNA Helicase and ATPase | yes | |
| UPF2 | UPF2 | UPF2, RNA Helicase and ATPase | ||
| UPF3B | UPF3B | UPF3B, Regulator of Nonsense Mediated mRNA Decay | ||
| USP10 | USP10 | Ubiquitin Specific Peptidase 10 | ||
| USP11 | USP11 | Ubiquitin Specific Peptidase 11 | ||
| USP13 | USP13 | Ubiquitin Specific Peptidase 13 | ||
| USP5 | USP5 | Ubiquitin carboxyl-terminal hydrolase 5 | ||
| USP9X | USP9X | Ubiquitin Specific Peptidase 9, X-Linked | ||
| UTP18 | UTP18 | UTP18, Small Subunit Processome Component | ||
| VASP | VASP | Vasodilator-stimulated phosphoprotein | ||
| VBP1 | VBP1 | VHL Binding Protein 1 | ||
| VCP | VCP | Valosin Containing Protein | ||
| WBP2 | WBP2 | WW Domain Binding Protein 2 | ||
| WDR47 | WDR47 | WD Repeat Domain 47 | ||
| WDR62 | WDR62 | WD Repeat Domain 62 | ||
| XPO1 | XPO1/CRM1 | Exportin 1 | ||
| XRN1 | XRN1 | 5'-3' Exoribonuclease 1 | yes | |
| XRN2 | XRN2 | 5'-3' Exoribonuclease 2 | ||
| YARS | YARS | Tyrosine—tRNA ligase, cytoplasmic | ||
| YBX1 | YB-1 | Y-Box Binding Protein 1 | ||
| YBX3 | YBX3/ZONAB | Y-box-binding protein 3 | ||
| YES1 | YES1 | Tyrosine-protein kinase Yes | ||
| YLPM1 | YLPM1 | YLP Motif Containing 1 | ||
| YTHDF1 | YTHDF1 | YTH domain family protein 1 | ||
| YTHDF2 | YTHDF2 | YTH domain family protein 2 | yes | |
| YTHDF3 | YTHDF3 | YTH domain family protein 3 | ||
| YWHAB | 14-3-3 | Tyrosine 3-Monooxygenase/Tryptophan 5-Monooxygenase Activation Protein Beta | ||
| YWHAH | 14-3-3 | 14-3-3 protein eta | ||
| YWHAQ | 14-3-3 | 14-3-3 protein theta | ||
| ZBP1 | ZBP1 | Z-DNA Binding Protein 1 | ||
| ZCCHC11 | ZCCHC11 | Zinc finger CCCH domain-containing protein 11 | ||
| ZCCHC14 | ZCCHC14 | Zinc finger CCCH domain-containing protein 14 | ||
| ZC3H11A | ZC3H11A | Zinc finger CCCH domain-containing protein 11a | ||
| ZC3H14 | ZC3H14 | Zinc finger CCCH domain-containing protein 14 | ||
| ZCCHC2 | ZCCHC2 | Zinc finger CCCH domain-containing protein 2 | ||
| ZCCHC3 | ZCCHC3 | Zinc finger CCCH domain-containing protein 3 | ||
| ZC3H7A | ZC3H7A | Zinc finger CCCH domain-containing protein 7A | ||
| ZC3H7B | ZC3H7B | Zinc finger CCCH domain-containing protein 7B | ||
| ZC3HAV1 | PARP-13.1/PARP-13.2/ARTD13 | Zinc Finger CCCH-Type Containing, Antiviral 1 | yes | |
| ZFAND1 | ZFAND1 | Zinc Finger AN1-Type Containing 1 | ||
| ZFP36 | TTP/TIS11 | ZFP36 Ring Finger Protein/Trisetrapolin | yes | |
| ZNF598 | ZNF598 | Zinc finger protein 598 | ||
| ZNF638 | ZNF638 | Zinc finger protein 638 |